Snowblind in the Desert Southwest: Moisture Islands, Ungulate Ecology, and Alternative Prehistoric Overwintering Strategies

Snowblind in the Desert Southwest: Moisture Islands, Ungulate Ecology, and AlternativePrehistoric Overwintering StrategiesAuthor(s): Alan J. OsbornSource: Journal of Anthropological Research, Vol. 49, No. 2 (Summer, 1993), pp. 135-164Published by: University of New MexicoStable URL: http://www.jstor.org/stable/3630187 .

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SNOWBLIND IN THE DESERT SOUTHWEST: MOISTURE ISLANDS, UNGULATE ECOLOGY,

AND ALTERNATIVE PREHISTORIC OVERWINTERING STRATEGIES

Alan J. Osborn

Department of Anthropology, University of Nebraska-Lincoln, Lincoln, NE 68588-0368

Archaeologists concerned with human adaptations in the American Southwest have generally assumed that plant resources dominated the diets of hunter-gatherers and cultivators throughout most of the prehistoric record. Such a perspective has probably arisen as a result of the "tyranny of the ethnographic record," the dominant role of "lowland archaeology," and the lack of research that was guided by robust ecologically based theoty. Optimal foraging theory, ungulate ecology, and animal and human physiology and nutrition suggest that upland areas, or "moisture islands," played a very significant role in the long-term evolutionary development ofArchaic and Anasazi-Hohokam-Mogollon populations throughout this region. Extant ecological knowledge of upland faunal populations, e.g., mule deer, elk, and bighorn sheep, and of their overwintering strategies is essential for our overall understanding of prehistoric life in this region.

IN THE AMERICAN SOUTHWEST, archaeologists have frequently assumed that diets and land-use strategies of early Archaic hunter-gatherers, late Archaic cultivators, and Anasazi-Mogollon-Hohokam agriculturalists were dominated by wild or domesticated plants (e.g., Cordell and Upham 1984; Euler et al. 1979; Ford 1984; Gasser and Kwiatkowski 1991; Leonard 1989; Minnis 1985, 1989; Fagan 1991; Sullivan 1987; Wills 1988, 1991, 1992; Winter 1980; Winter and Wylie 1974; Winter and Hogan 1986). Archaeologists have often relied on the empirical generalizations of Julian Steward (1938), i.e., the "Great Basin Shoshonean model," and Jesse Jennings (1957, 1964, 1978), i.e., the "Great Basin Desert Culture," to reconstruct Archaic hunter-gatherer lifeways in the American Southwest (Frisbie 1983; Reed 1964; Schaafsma 1983). One of the most powerful challenges to such views of western Archaic adaptations comes from optimal foraging theory. Diet choice models, for example, predict that plant foods such as grass seeds, cattail pollen, pinyon nuts, and tubers were added to human diets relatively late in the prehistoric period due to their high handling costs (O'Connell, Jones, and Simms 1982; Simms 1984, 1985, 1987). Considerable archaeological evidence also indicates greater prehistoric dependence on bighorn sheep in the Great Basin and the Colorado Plateau than was observed by Euro-Americans (Pippin 1977). As a consequence, historic accounts of Great Basin hunter-gatherer diets provide an inappropriate repre- sentation of earlier food-getting strategies.

(Journal of Anthropological Research, vol. 49, 1993)

135



136 JOURNAL OF ANTHROPOLOGICAL RESEARCH

Archaeological interpretations of Anasazi, Fremont, Mogollon, and Hohokam adaptations have, until recently, minimized the role of hunting. Instead, a plethora of studies has focused on the significance of maize horticulture and growing season climatic factors (e.g., Dean et al. 1985; Euler et al. 1979; Haury 1976; Hill 1970; Holbrock and Mackey 1976; Jorde 1977; Karlstrom, Gumerman, and Euler 1976; Lindsay 1986; Moore and Winter 1980; Petersen 1988; Plog 1979; Schoenwetter and Dittert 1968; Schlanger 1988; Vivian 1970). Many reconstructions of prehistoric agricultural lifeways were based on overly simplified Hopi, Zuni, Pueblo, or Pima analogues (cf. Fish and Nabhan 1991; Gasser and Kwiatkowski 1991; Hunter-Anderson 1986; Martin 1985; Powell 1983; Sullivan 1987).

Ethnographic, as well as geographic, studies in the American Southwest frequently stressed the causal connections between maize farming, spring and summer precipitation, population movements, and other aspects of aboriginal life in this region (Beaglehole 1936; Bradfield 1971; Dozier 1970; Eggan 1950; Forde 1931; Hack 1942).

This paper is a reassessment of the role of hunting in the long-term evolutionary development of prehistoric societies in the American Southwest.' Optimal foraging theory and derivative models predict that both wild plant exploitation and domesticated plant production in this region might best be understood as a function of the decreased availability of higher-ranked food resources, i.e., large mammals including bighorn sheep, elk, mule deer, bison, and antelope. These large ungulates were restricted to the more mesic, upland communities that developed during the early Middle Holocene (ca. 8000 B.P.; Van Devender and Spaulding 1979).

We can expect that these ungulates played a significant role in the adaptations of past foraging, collecting, and cultivating societies in, and adjacent to, upland areas of the American Southwest (Figure 1). Consequently, considerable attention will be given in the first portion of this paper to environmental factors that condition the distribution, abundance, accessibility, and physical condition of ungulates-particularly in winter. Such a reassessment of prehistoric hunting in this region must entail explanatory models that systematically link climate, physiography, ungulate ecology, and human adaptations. These models may prove to be relatively radical departures from previous archaeological research in the American Southwest. A comprehensive understanding of prehistoric adaptations during the Archaic, Basketmaker, and portions of the Anasazi- Mogollon-Hohokam periods must incorporate environmental variables that con- trolled the abundance, distribution, and condition of ungulates in highland areas. We must turn our attention to winter precipitation and associated climatic factors throughout the Colorado Plateau including the La Sal, Abajo, Henry, Chuska, San Juan, Sangre de Cristo, Sacramento, Black Range, Mogollon, and Santa Catalina mountains and the Wasatch, Uncompahgre, Kaiparowits, and Kaibab plateaus. An explanatory model based on winter climate, ungulate ecology, and prehistoric hunting will establish the initial conditions for the later adoption of food storage and agriculture in the American Southwest.



SNOWBLIND IN THE DESERT SOUTHWEST 137

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ABORIGINAL HUNTING IN THE AMERICAN SOUTHWEST

Aboriginal hunting in the American Southwest has been discussed in a number of archaeological and ethnographic contexts (e.g., Beaglehole 1936; Cordell 1977; Davis 1960; Emslie 1983; Hastorf 1980; W.W. Hill 1938; James 1990; Neusius 1984; Neusius and Phagan 1983; Powell 1983; Sharp 1989, 1990; Speth 1991; Speth and Scott 1985, 1989; Szuter 1991; Szuter and Bayham 1989; Tainter 1984; Wetterstrom 1986; Young 1980). Most studies have assumed that both wild and domesticated plants comprised the bulk of the diet and that ungulates and other animals were a supplemental food resource.

Some investigators have recently begun to assess the role that animal resources played in local and regional-level feeding strategies, food exchange systems, and the long-term evolution of aboriginal societies in the American




Southwest (Speth 1991; Speth and Scott 1985, 1989; Spielmann, Schoeninger, and Moore 1990; Szuter 1991; Szuter and Bayham 1989).

Any assessment of the role of large mammals in prehistoric diet must consider the differential value of animal flesh and fat as a source of energy; essential amino and fatty acids; fat soluble vitamins A, D, E, and K; iron; and zinc. In general, the caloric utility of animal resources can be expected to vary inversely with levels of plant consumption. Animal resources will be used primarily as a nutrient source once carbohydrate and oil-rich plant foods begin to provide the greatest proportion of the food energy demands.

Hunter-Anderson (1986:33) argued that mountain ranges in the American Southwest exhibited higher primary productivity, and, therefore, hunter-gatherer populations were more dense in these areas. She (1986:36) proposed that "the overwintering problem" posed by a shortened growing season in the higher elevations was resolved by the adoption of plant storage. The significance of large mammals is downplayed since it is assumed that ungulates were dispersed throughout the year and that meat preservation would have been difficult in this region.

Szuter and Bayham (1989) found that hunting throughout the Archaic and Hohokam periods (2500 B.C.-A.D. 1350) in south-central Arizona did not de- crease in importance. With increased regional population density and reduced residential mobility, lowland Hohokam settlements would have established hunting camps in the uplands for the procurement of higher-ranked artiodactyls. Animal products were presumably transported back to lowland residential sites during the Hohokam period (A.D. 500-1350). For the Hohokam, winter and spring food shortages would have been reduced through use of stored agricultural crops (Szuter and Bayham 1989:87). Little discussion is given to environmental factors that condition wild animal distribution, accessibility, and abundance- particularly for artiodactyls in this region.

Speth and Scott (1985, 1989) discussed archaeological evidence for an increased dependence on large mammals (i.e., deer, mountain sheep, antelope, and bison) that included analyses of Ventana Cave (Hohokam components), Snaketown, and Point of Pines in Arizona and of Gran Quivira, Pecos Valley sites, Arroyo Hondo, Chaco Canyon, and the McKinley Mine area in New Mexico (Speth and Scott 1989:72-73). Speth and Scott (1989:73) stated,

In sum, the widespread shifts in the Southwest toward progressively greater reliance on large mammals, especially deer, antelope, mountain sheep, and bison, do not seem to be direct responses to altered climatic or environmental conditions, changes in the absolute size of regional populations, or technological innovations.

The high-quality protein demands of prehistoric horticultural settlements were met by sending task groups to more distant hunting areas or by ex- changing plant food products for meat (Speth and Scott 1989:78). Speth and




Scott (1989) did not consider the impact of local and regional climatic conditions on large mammals such as mule deer, elk, and bighorn sheep in the upland areas.

ECOLOGICAL BACKGROUND

Physiography, Flora, and Fauna Mesic environments that supported sizable populations of ungulates existed

as "isolated islands of moist montane habitat in a sea of desert" (Erlich, Murphy, and Wilcox 1988). These mountains, plateaus, and questas in the Basin and Range and Colorado Plateau provinces have been referred to by ecologists and meteorologists as "rainy islands" (Bailey 1981), "alpine islands," or "mountain islands" (Bender 1982).

These higher-elevation settings exhibit cooler temperatures, increased precipitation, and decreased evaporation (Pianka 1983). Arid lands throughout the Colorado Plateau are referred to as "cool deserts" because they "are located in higher latitudes and stand at higher altitudes than is typical of many other arid regions of the world" (Bailey 1981:13). These "moisture islands" result from orographic effects that increase both winter and summer precipitation.

These highland areas are characterized by a series of parallel or concentrically arranged and heavily vegetated zones including pinyon-juniper (1,524-2,134 m), mountain brush (1,981-2,438 m), montane fir-aspen (2,042-2,743 m), and subalpine spruce-fir (2,850-3,450 m) communities. Primary productivity associated with these moisture islands is substantially greater than that of the surrounding shadscale, big sagebrush, and blackbrush communities that occur below 1,500 m. Winter forage in the La Sal Mountains of southeastern Utah ranges from 40 dry grams/m2 in the desert shrub community to 240 dry grams/ m2 in the mountain brush and grass community (Pederson 1970:56, table 13). These moisture islands sustain relatively large resident populations of large ungulates, including Rocky Mountain mule deer (Odocoileus hemionus), elk (Cervus canadensis), and bighorn sheep (Ovis canadensis).

Ungulate Physiology and Nutrition Numerous wildlife ecology studies demonstrate that North American un-

gulates (e.g., mule deer, bighorn sheep, and elk) living in high-elevation or high-latitude settings exhibit marked fluctuations in body composition, particularly fat, throughout an annual cycle (e.g., Anderson, Medin, and Bowden 1972; Cheatum 1949; Fong 1981; Franzmann and Arneson 1976; Hunt 1979; Kie 1978; Medin and Anderson 1979; Miera and Schmidt 1981; Ransom 1965; Snider 1980). Anderson, Medin, and Bowden (1972) and Miera and Schmidt (1981) have conducted detailed analyses of the seasonal fluctuations in carcass, back, kidney, and bone marrow fat for Colorado mule deer (Figures 2 and 3). The bulk of body fat is deposited during summer and early autumn because of the abundance and high nutritional value of forage then. Deer exhibit their best




100 Male Carcass Fat

90 '"" '."' "'"'.-

" . . ,..

.. .. ... .. . . ... . Male

0 Femur Fat

Female 70 ................................. Carcass Fat

6 0 ........................................ ................................. .................. F em a le Femur Fat

50...................

30.............

20 .............

10

0 I I Autumn Winter Spring Summer

Season

Figure 2. Seasonal Changes in Carcass Fat and Femur Marrow Fat for Mule Deer in the Cache la Poudre River Drainage, Colorado

Based on data from Anderson, Medin, and Bowden (1972:586, table 2).

condition during October, just prior to the breeding season. Peak predicted weight losses for mule deer are 17.2 kg (37.9 lbs), or 19 percent, for males and 16.0 kg (35.2 lbs), or 22 percent, for females (Anderson 1981:72). Carcass fat in male mule deer peaks in autumn and declines sharply through winter until early spring; does exhibit minimum fat levels in late spring/early summer (Anderson, Medin, and Bowden 1972 quoted in Short 1981:123).

Mule deer do not increase food consumption levels during winter but rely, instead, on fat reserves. Body fat deposits do not appear to provide much insulation during the winter. As Anderson (1981:74) states, "body maintenance, not insulation, probably was the primary function of subcutaneous fat in that population at 40 degrees 40 minutes north latitude." This body fat is used to supplement lower forage intake during the winter in this region of North Amer- ica. Harris (1945) suggested that fat deposits are used in the sequence of subcutaneous, visceral, and femur marrow fat (Anderson 1981:71).




100 Males-CP

90 E Males- M 90-------------------------------M-esM

FemalesMV

! 70 - ---------------------- - -

60 \ - --- --

50-----------------------------

50

2n 0 ----'--- -------- - -

30 ------------ 20

Autumn Winter Spring Fall

Season

Figure 3. Seasonal Changes in Kidney Fat for Mesa Verde (MV) and Cache la Poudre (CP) Mule Deer in Colorado

Based on data from Miera and Schmidt (1981:19, table 3).

Aspects of Ungulate Overwintering The most severe environmental constraints imposed on the ungulates of

North America, and the Northern Hemisphere in general, coincide generally with the onset of winter (Edwards 1956:159; Formozov 1946; Nasimovich 1955). The significance of this stress period has been discussed for white- tailed deer (Crete 1976; Spiker 1933), black-tailed or mule deer (Jones 1975; Strickland and Diem 1975), pronghorn antelope (Rand 1947), Dall sheep and moose (des Meules 1964; Murie 1944), the American bison (Daubenmire 1985), mountain goats (Cowan 1950, 1952), elk (Banfield 1949; Mathews 1952), and reindeer or caribou (Pruitt 1959; Wildhagen 1952). Severinghaus (1947) first showed that deer mortality in the Adirondacks of New York "varies from year to year with the severity of the weather" and that "deep snow is the principal weather factor involved" (Edwards 1956:159).

Ungulates' Abilities to Cope with Snow. Telfer and Kelsall (1984) have rank ordered the ability of a number of North American mammals to cope with snow (Table 1). They developed a morphological index (MI = MCH + [100 + (MFL/10)]), where MCH = mean chest height (cm) and MFL = mean foot




loading (g/cm2). They also used behavioral characteristics to assess the winter coping abilities of mammals. These characteristics include ability to feed on aboveground food, use of trails, rooting/digging ability, migration (horizontal or vertical), selection of most suitable winter ranges, and techniques of locomotion (Telfer and Kelsall 1984:1829). Winter snow severity was based on depth, as well as crust characteristics. They (1984:1831) recognized that there was a "regular increase in [the] mean adaptation index . .. for regional faunal groupings from the short-grass plains species to the boreal forest [that] corresponds ... to the ... severity of snow conditions."

Although white-tailed deer and mule deer exhibit relatively low morphological indices for snow coping ability, they can be found in areas that exhibit severe winter conditions. They appear to be able to respond to these conditions via increased behavioral flexibility (see Table 1).

The snow coping abilities of North American ungulates may vary with respect to sexual dimorphism. Different snow coping abilities related to morphology and sex may have behavioral implications. According to Telfer and Kelsall (1984:1832),

Moose show the least sexual dimorphism, which perhaps accounts for the few observations of sexual aggregation on winter ranges used by moose. White-tailed deer exhibit some sexual dimorphism, which should give males a slight advantage in deep snow. Laramie and White (1966) reported that adult male deer often wintered separately from other deer, and were in deeper snow farther up the mountain sides. .... However, female deer have a slightly lower foot loading, which would give them a survival advantage on crust or dense snow.... In addition, female and juvenile deer overwinter in large aggregations, and their more extensive trail network may have provided greater mobility and more access to forage during the period of thick snow cover.

TABLE 1 Snow Coping Abilities for North American Ungulates

Mean Snow Morphological Behavioral Coping

Species Index/200 Index/30 Index

Caribou .77 .87 .82 Moose .70 .63 .67 Dall sheep .61 .57 .59 Wapiti .59 .60 .60 Bighorn sheep .57 .53 .55 White-tailed deer .56 .70 .63 Bison .48 .50 .49 Antelope .41 .43 .42

Source: Telfer and Kelsall (1984:1831, table 3).




Parker, Robbins, and Hanley (1984:481) found that the energetic costs of locomotion in snow for mule deer and elk are dictated by travel velocity, sinking depth, and snow density. They (1984:481) found that "the net energy cost of travel for a 100-kg elk calf in 59 cm of snow is approximately five times the cost of locomotion without snow." Locomotion costs (kcal/kg/min) for elk increased exponentially relative to sinking depth (rY = .98) and increased linearly relative to travel velocity (r2 = .94-.96; Parker, Robbins, and Hanley 1984:482).

Snow Cover and Winter Range Conditions. Snowfalls in fall, winter, and spring trigger ungulate movements from high-elevation summer ranges to lower-elevation winter ranges. Rocky Mountain mule deer begin to move from higher-elevation summer range to intermediate-elevation winter range once snow accumulation equals 10-25 cm (8-10 in). This downward migration generally starts in late October (Gilbert, Wallmo, and Gill 1970:16). Mule deer descend to lower elevations in increasing numbers as snow begins to accu- mulate at the higher elevations. Very few mule deer were observed in areas where the snow accumulation exceeded 60 cm (24 in). These ungulates tend to range at the highest elevations that winter snows allow (Gilbert, Wallmo, and Gill 1970:18).

Mule deer make more limited use of the lower elevations in arid regions because scant summer rainfall, high temperatures, and high evaporation greatly limit forage production during the growing season (Wallmo 1981:16-17). The return to higher-elevation summer range may be delayed or reversed if temperatures drop, wind velocity increases, and snowfall occurs during the spring migration (Richens 1967:656).

Ungulates frequently make use of microenvironmental variation and snow cover to minimize cold stress and foraging costs (Robinette et al. 1952:290). Richens (1967:664) provides a succinct discussion of mule deer winter foraging areas and microenvironmental preferences:

Local deer numbers were determined by availability of food, snow depth and condition, temperature, wind, and the amount of protective cover. In cold, windy, or stormy weather deer were usually found in sheltered areas, particularly in heavy juniper stands. On cold, sunny days most deer occupied bare southern slopes where temperatures were highest and snow depth the least. Northern slopes were used heavily in the fall and spring but were abandoned during the winter when they were covered with deep, crusted snow.

Geist (1971:92) described the movements of mountain sheep in response to fall and winter snowfalls. In early winter, sheep sought grassy slopes with deep, soft snow cover or windswept talus slopes with southern exposure. Geist (1971:94) stated, "The wintering sheep, ibex, or mountain goats move where forage is most easily available, where the snow has been removed by wind, sunlight, avalanches, or gravity, or where little snow fell on the ground in the first place." In late winter and early spring, mountain sheep move to




south-facing cliffs and boulder fields where snowmelt is heaviest and forage is accessible.

Winter Range Size and Ungulate Densities. Ungulates frequently reduce their range size during the winter months. For example, mean winter range size for white-tailed deer in southern New Brunswick was strongly and inversely related to snow depth (r = -.94; p < .02; Drolet 1976:125). During severe winters, mule deer on the northern slopes of Utah's Uinta Range "occupied only 60.5 percent of the area used during the normal winters, with upper range limits at 6,500-7,000 ft. elevation. ..." (Richens 1967:664). Mule deer density in this region ranged from 81 to 135 deer per square mile during the winter, whereas the average summer and winter density equaled 13 deer per square mile (Richens 1967:664). Winter deer counts in north-central Col- orado were also shown to be inversely related to mid-to-late-winter precipitation (Gilbert, Wallmo, and Gill 1970:21).

Winter Severity and Ungulate Fertility. Mech et al. (1987) have demon- strated a set of relationships between winter snowfall, body weight, and the fertility of white-tailed deer and moose populations. These ungulates experi- ence a pronounced period of weight gain during the spring, summer, and fall; considerable weight is then lost throughout the winter. The degree of weight loss is a function of the length and the severity of winter. Winter severity involves duration of freezing and below-freezing temperatures, the depth and duration of snow, and the extent of "crusting" or ice formation. Crust formation due to melting and freezing impedes the movements of ungulates and limits access to adequate forage. These climatic conditions, then, ultimately determine the extent of the annual weight loss and the ability of the animals to recover during the rest of the year.

Deer and moose fertility depends considerably on the degree of winter malnutrition and warm-season recovery. Furthermore, females are pregnant during winter and spring. For this reason, winter and spring weather potentially has an enormous effect on female weight loss, fetal development, survivability of offspring, and ultimately the degree of annual population growth (Mech et al. 1987:616). Gilbert, Wallmo, and Gill (1970) found that the percent of in- terannual change in mule deer population size was inversely related to winter precipitation expressed as inches of water (r = -.92) or expressed as inches of snow (r = -.79).

Mech et al. (1987) hypothesized that snow accumulations in previous, con- secutive winters would have an inverse, cumulative effect on future deer and moose productivity and population changes. These investigators used linear regression analyses to demonstrate a causal relationship between previous winter climatic conditions and changes in ungulate population. The results of these analyses are presented in Table 2. Fawn-to-doe ratios for mule deer and lamb-to-ewe ratios for bighorn sheep have also been shown by Picton (1984) to be inversely related to late winter and early spring snow depth and water content (Table 3).




TABLE 2 Relationship between Sum of Previous Winters' Snow Accumulation and

Measures of Population Productivity in Deer and Moose

Species Dependent Variables r R p

White-tailed deer Fawn-to-doe ratio .60 .36 <.01 Moose Calf-to-cow ratio .26 .51 <.01 Moose Twinning rate .18 .42 <.01 White-tailed deer Population change (%) .20 .45 <.01 Moosea Wolf predation rate .24 .49

on calves (8-10 mos)

Source: Data are from Mech et al. (1987:620, table 3). a. Regression results for moose are based on previous 5-7 years of snow accumulation.

Winter Severity and Ungulate Mortality. Picton (1979:117) found that winter climatic conditions accounted for approximately 70 percent of the variation in fawn survival. Warm, dry winters favored increased production and survival of fawns. Climatic conditions determine the nutritional status and reproductive capabilities of female ungulates. Short (1981:121) found: (1) well-nourished female mule deer lost 5 percent of their fawns; (2) winter-stressed does lost 33 percent; and (3) does on deficient diets throughout gestation lost 90 percent of their offspring.

Hobbs (1989) developed a simulation model of energy balance for mule deer does and their fawns. Mortality increased from 1.3 percent to 22.7 percent for does and from 4.2 percent to 72.1 percent for fawns as winter climate

TABLE 3 Correlations between Winter and Spring Precipitation and Offspring

per Female for Mule Deer and Bighorn Sheep in the Sun River Drainage, Montana

Species Independent Variable r R p Mule deer March 1 snow pack

water content .545 .30 >.05 April 1 snow depth .564 .32 >.05

Bighorn sheep March 1 snow pack water content .92 .85 >.05

March 1 snow depth .96 .92 >.01 April 1 snow pack

water content/snow depth .88 .77 >.05

Source: Data are from Picton (1984:874, table 1).




increased from mild to severe. Higher mortality rates were related more to decreased forage availability than to increased energy costs for winter activity.

Snow Cover and Predation. Winter severity can also be related systematically to variable levels of predation on ungulate populations (Peterson and Allen 1977; Nelson and Mech 1981, 1986). Nelson and Mech (1986:471) state, "Wolves capture more prey during severe winters with deep snow." These investigators (1986:472) found "a significant positive relationship between Jan- uary-April wolf predation rates and January-April snow indexes, which explained 51% of the variation in predation rate."

Snow significantly affects the escape capabilities of deer. It restricts mobility, increases energy costs, and reduces deer fat reserves (Mattfeld 1974; Parker, Robbins, and Hanley 1984). Cold temperatures also deplete deer fat reserves because of increased maintenance costs resulting from heat losses. "The cumulative effect of this energy drain, especially in late winter, decreases deer physical condition and predisposes them to wolf predation" (Nelson and Mech 1986:472).

ALTERNATIVE MODEL FOR PREHISTORIC OVERWINTERING STRATEGY

The primary portion of this explanatory model (Figure 4) is applicable to large mammal hunting for foragers, collectors, and cultivators in the Colorado Plateau and other mountainous areas of southern New Mexico and Arizona. This ungulate procurement model can also be modified and extended to account for prehistoric overwintering strategies in portions of the northwestern Plains, the Great Basin, California, and northern Mexico. It can be used to explain considerable variation in land-use and mobility strategies, diet breadth, food storage, adoption and use of ceramic vessels, human demography, and sedentism.

The previous discussion delineated some of the most significant causal relationships that link winter precipitation to ungulate mobility, home range size, fertility, mortality, diet, and body composition. The determinant linkages between snowfall and ungulate ecology serve as the basis for the construction of explanatory models for aboriginal hunting strategies in the American South- west. Excessive winter snowfall, low temperatures, and high winds in upland areas of the American Southwest greatly limit ungulate mobility, availability of high-quality forage, body fat reserves, thermoregulation capabilities, and offspring survival. At the other extreme, scant snowfall in these mountainous regions delays the aggregation of ungulates and their migration to winter ranges at lower elevations. Limited snowfall may also mean that mule deer will continue to make use of relatively large home ranges during the winter. Scant snow cover would also mean that deer, elk, and bighorn sheep would be less apt to move to smaller, more predictable areas within their winter range. Limited winter moisture would also reduce forage productivity and lower its nutritional value during the next growing season. Inadequate quantity and quality of forage




HIGH ELEVATION OROGRAPHIC SETTINGS

PRECIPITATIONADOPTION/USE

ADOPTION-USE

CERAMIC VESSELS

GROWING WINTER PLANT SEASON DETOXIFICATION

RAINFALL & PROCESSING

FORAGE UNGULATE USE OF PRODUCTION SNOW COVER -- PREDATION "GRANARIES"

RATES

FORAGE UNGULATE

ONSLIMITED QUALITY & SNOW COPING

DOWNSLOPE WILD PLANT ACCESSIBILITY ABILITIES

MIGRATION STORAGE

UNGULATE UNGULATE REDUCED LIMITED CONSUMPTION METABOLIC HOME RANGE WILD PLANT

COSTS SIZE CONSUMPTION

UNGULATE UNGULATE DENSITY ETHERMOGENIC

CONDITION BODY FAT PREDICTABILITY FOOD COSTS

UNGULATE MENATEVALUE MORTALITY FAT/PROTEINUNGULATE

RATIO HUNTING

UNGULATE FERTILITY POPULATION

Figure 4. Generalized Model for Prehistoric Ungulate Procurement in the American Southwest

will retard the accumulation of body fat reserves. In turn, low fat reserves will reduce the capabilities of ungulates to endure more severe winters in the future.

Petersen (1988) has made a significant contribution to monitoring the long- term variation in winter precipitation in this region. Patterns in winter precipitation can then be causally linked to the preceding relationships between climate and ungulate ecology. For example, Dix and Richards (1976) found that spruce trees exhibit a low heat and drought tolerance. The distribution of spruce throughout the Rocky Mountains corresponds to areas of long-term snowpack and lack of summer drought conditions (Dix and Richards 1976:311). Petersen (1988) remarked that "The actual advancement of spruce trees downslope depends on the successful establishment and survival of seedlings. Seeds are set in the fall and spend the winter under the snow." He proposed that spruce-to-pine pollen ratios can be used to monitor long-term shifts in winter versus summer dominant precipitation. Petersen (1988:82) found that spruce/ pine pollen ratios greater than 0.60 reflect winter precipitation like that of the




present-day regime and ratios less than 0.60 indicate less winter precipitation than occurs today. Spruce/pine pollen ratios have been calculated for the time period 900 B.C. to the present in the La Plata Mountains of southwestern Colorado. This ratio exceeded present-day values for the periods 100 B.C.- A.D. 500, A.D. 600-800, and A.D. 1000-1100 (Petersen 1988:75, fig. 40). Such climatological data for winter precipitation could be used to model long-term fluctuations in snow cover and ungulate populations in this region.

High-elevation questas and mesas like Mesa Verde and Black Mesa, plateaus such as the Kaiparowits and Kaibab areas, and mountain ranges including the La Platas, La Sals, Abajos, and Chuskas supported sizable populations of mule deer, elk, and bighorn sheep. Isolated peninsulas, islands, and archipelagoes formed by mountain systems would have been preferred resource patches for human hunters in this region-particularly during the winter. Archaeologists have generally underestimated the resource potential of such large mammals (cf. Grady 1980).

The present-day mule deer population of Arizona, New Mexico, Colorado, and Utah can exceed 1.34 million animals. Maximum harvests by modern hunters in this four-state region produced 366, 000 animals in one year (Connolly 1981). The flesh and fat of an adult mule deer yields approximately 71,268 kcal. This maximum harvest of mule deer could have supported 31,000 persons for eight months (given that an adult uses 3,500 kcal/day for 240 days). In the past several decades, the deer population in this region has declined to a minimum of 722,000 animals (Connolly 1981). Mule deer harvest at that time equaled 107,000 animals. This minimal harvest could have supported approximately 9,046 individuals for about eight months. Such gross estimates of prehistoric hunter-gatherer populations do not include elk and bighorn sheep. Calculations involving contemporary mule deer populations also ignore the effects of livestock overgrazing, fire prevention, predator control, and development projects.

Ungulates would have been a significant calorie source in winter for prehistoric hunter-gatherers in upland areas of the American Southwest. Deer, bighorn sheep, and elk exhibit higher return rates (kilocalories per unit handling time) relative to plant food resources (Simms 1984, 1987; Figure 5). Food energy is the most relevant currency in this case since it is a limiting factor in nutritional, physiological, and ecological processes. Stored plant foods would then be added to hunter-gatherer diets in response to the decreased availability of these high-return animal resources (O'Connell, Jones, and Simms 1982; Simms 1984, 1985, 1987). Plants require considerable handling costs with respect to food processing (e.g., hulling, grinding, roasting, winnowing, baking, and boiling; Hawkes and O'Connell 1981). They may also represent considerable metabolic costs owing to the presence of toxins and antinutrients, such as alkaloids, polyphenols, lectins, phytates, oxalates, proteinase inhibitors, and nonprotein amino acids (Bressani, Elias, and Braham 1982; Johns 1989; Johns and Kubo 1988; Rosenthal and Janzen 1979; Stahl 1989; Thomas, Samson, and Bergeron 1988; Walker 1982).




35

*Minimum

SMaximum

25 ?-

5 ?

5 - ---

-

-el oP Q~ Is "t~4e .?e?

Figure 5. Return Rates for Select Food Resources in Western North America

Based on data from Simms (1984:93, table 5).

Both female and male mule deer undergo seasonal changes in body composition that specifically involve fat levels. Thus, for human hunters, the energy returns from the same unit weight of food decline markedly from summer and fall through the winter and spring (Figure 6). Ungulates exhibit significant declines in energy value due to shifts in the ratio of fat-to-lean carcass weight.

Increased protein-derived energy relative to fat-based energy is associated with increased diet-induced thermogenesis (Jequier 1983; Woo, Daniels-Kush, and Horton 1985). Diet-induced thermogenesis (DIT) varies with respect to the composition of one's diet: generally, DIT equals 4 percent for fat, 6 percent for carbohydrates, and 25-30 percent for protein. In other words, 20-30 calories of protein would be lost as heat following ingestion of 100 calories of protein. If ingested simultaneously, carbohydrates and/or fats can reduce the higher costs of protein metabolism. Carbohydrates exhibit a greater protein- sparing effect than fats; this effect is quite pronounced for individuals with less than optimal caloric intake (Speth 1983:154). Speth (1983:155) also points out that "the negative nitrogen balance produced by fat substitution may persist for much longer periods and may have a much more detrimental effect." During




90 Adult Female

Adult Male 80 -- - - - -----------------

70 ----------------------------------

50 ------ -----------------------

80

30 -- -- -------- - 20

Autumn Wimter Spring Summer

Season

Figure 6. Seasonal Changes in the Food Energy Value of Mule Deer Protein and Fat

Based on data from Anderson, Medin, and Bowden (1972:586, table 2).

seasonal lows in resource availability and prey condition, prehistoric hunter- gatherers lacking suitable carbohydrate foods preferentially processed and consumed animals that provided the highest fat content in order to offset the inordinately high caloric costs of protein metabolism (Speth 1983, 1987, 1990, 1991; Speth and Spielmann 1982).

Consequently, the energetic costs for aboriginal hunter-gatherers overwintering in these higher-elevation settings would have been compounded. The energy value of ungulates undergoes a marked reduction throughout the winter and spring. At the same time, a meat-based diet becomes more and more costly with respect to metabolic costs related to increased diet-induced thermogenesis. This increased cost may rise to approximately 30 percent of resting metabolic rate (RMR). Additional stress would also be imposed by cold temperatures and a corresponding increase in adaptive thermogenesis (AT), which involves physiological responses to cold stress in this case (Woo, Daniels- Kush, and Horton 1985).

Hunter-gatherers in the mountainous areas of western North America most probably adopted an overwintering strategy that initially minimized their commitment to food storage. As noted, ungulates are relatively well adapted to the rigors of winter throughout western North America. Mule deer, elk, and




bighorn sheep exhibit relatively high snow coping abilities related to morphology, physiology, and behavior. These animals have essentially solved the overwintering problem (Binford 1980). Human populations, in turn, could have relied on stalking and ambush hunting throughout the winter. For example, the Ute of the southern Rocky Mountains wore snowshoes and hunted elk in deep snow; the animals were killed after they tired in the snow (Smith 1974:54 in Callaway, Janetski, and Stewart 1986:341). Snowshoes, in these cases, greatly reduced the ergonomic costs of traveling in deep snow. An archaeological example of Anasazi snowshoes is illustrated in Nordenskibld (1979:plate 48- 2).

If aboriginal groups overwintered in mountainous areas where ungulates were compressed within small winter ranges by deep snows, we might expect to observe winter camps located within or above the pinyon-juniper zone. As Callaway, Janetski, and Stewart (1986:343) note,

Fall gathering of pinon nuts was combined with deer hunting. .... Several families or even several bands might congregate and live mostly on pinon nuts and venison until deep snows forced a retreat to lower elevations. Pinon nut stores were revisited to carry supplies to camps at lower elevations (0. C. Stewart 1942:250).

Winter residential sites probably included relatively short-term camps in areas where game was less constrained by winter snowfall and was therefore less predictable. These low-visibility camps might be similar to those described by John Wesley Powell (1868-1880, in Fowler and Fowler 1971) for the Northern Ute in southern Utah and northern Arizona.

Other residential sites may have been more permanent or subject to re- petitive use throughout a number of years. These sites might look similar to those used by the Northern Paiute, who overwintered in the mountains and occupied substantial semisubterranean, conical winter houses constructed of juniper and pinyon poles, brush, and earth. Archaeological examples of such an overwintering strategy might look like Aspen Shelter in central Utah. This small rockshelter is located at 2,500 m (8,200 ft) above sea level. It contained two shallow basins that were interpreted as house structures; radiocarbon dates ranged from 4000 to 1300 B.P. (Janetski, Crosland, and Wilde 1991). The faunal assemblage was dominated by mule deer remains.

In order to circumvent diminishing energy returns and increased diet-induced thermogenic costs, aboriginal hunter-gatherers could have adopted a limited plant storage strategy based on dropseed (Sporabolus sp.), Indian rice grass (Oryzopsis sp.), peppergrass (Lepidium sp.), and giant wild rye (Elymus sp.), as well as oil-rich seeds and nuts such as wild sunflower (Helianthus sp.) and pinyon nuts (Pinus edulis). The bulk of these plant resources was generally obtained at lower elevations. These plant resources were visited and exploited in the spring, summer, and fall seasons during periods of peak availability. Hunter-gatherers would have exploited extensive home ranges during the




growing season. Caches of these resources were most probably made during the harvest close to productive patches. These caches could then have been visited throughout the winter by task groups that resided at higher elevations. With respect to the Southern Paiute, Kelly and Fowler (1986:371) state,

For most, fall was a time of plenty and one of great mobility, with shuttling from high to low country and from one spot to another where collecting and hunting were most favorable. Many moved to the mountains to cache pine nuts and hunt large game, returning to the valleys for rabbit drives. Highland seeds and berries were available, as was yucca fruit. All groups stored as much food as possible against the winter and recurrent spring famine.

Small isolated "granaries" that occur throughout much of southeastern Utah probably reflect this form of overwintering strategy (Figure 7). Although no systematic study has been conducted, many of these features do not appear to be associated with long-term residential locations. They have been observed throughout southeastern Utah in Canyonlands, Cedar Mesa, Glen Canyon, and elsewhere. These storage facilities were constructed of dry-laid masonry and wet-laid masonry within relatively small, narrow overhangs or crevices (Gun- nerson 1969:150).

The dry-laid masonry structures perhaps contained foodstuffs and water in sealed gourd or ceramic containers, as well as a variety of raw materials, implements, and facilities. The wet-laid, mud-plastered structures could have contained unprocessed foods in loose form or foodstuffs placed in vegetal fiber sacks, baskets, gourds, or ceramic vessels.

In addition, a number of isolated caches of ceramic vessels have been located throughout the American Southwest and in southern California (Figure 8). These ceramic vessel caches frequently contained foodstuffs such as roasted pinyon nuts, mescal cakes, and grass seeds. The vessels were frequently hermetically sealed with lac from creosote bushes (Euler and Jones 1956; Osborn 1991; Osborn and Wolley 1993). Both "granaries" and vessel caches could have been visited periodically by logistical groups from winter camps at higher elevations.

These plant foods yield from 1,400 to 4,800 kilocalories per kilogram. Jones and Madsen (1989) have recently calculated that conical carrying baskets used historically in the Great Basin could transport a minimum of 21,672 kcal of wild rye or a maximum of 171,623 kcal of pinyon seeds. One basket load of pinyon seeds would equal 2.4 mule deer (average carcass yield). The carbohydrate and oil-rich plant resources could then be consumed in limited quantities to offset the increased DIT costs of consuming ungulates during the winter.

Variation of winter severity in the uplands would have imposed significant constraints on this overwintering strategy. Scant winter snowfall would fail to force ungulates to migrate from higher-elevation settings in the fall, to use smaller winter home ranges, and to feed in restricted areas with southern




. .... . . .'

. .. i. ~ ?

. •

"; .

%

11

.r.

,, %,"ow •

0 jo, *; ss

Figure 7. Wet-Laid Masonry Storage Structure with Fitted Rock Slab Door at Big Ruin (42SA1586)

Courtesy Midwest Archeological Center, NPS

exposure. This would increase the costs of hunting, and hunter-gatherers might shift to increased dependence on plant storage in the ensuing winters. In addition, periods of severe winter conditions and deep snows might have dec- imated ungulate populations and forced increased dependence on stored plant resources for several years. Continued stress imposed by extremely dry winters or cold, wet winters would require that aboriginal hunter-gatherers adopt a more specialized storage strategy based on well-monitored patches of energy- rich, storable foods including domesticated plants. Ecological models about food storage, as well as its effects on human female fertility, have been discussed elsewhere (Osborn and Wolley 1993).

Once available, domesticated plants, including maize, beans, and cucurbits, may have been used initially to minimize the costs associated with hunting and




? ri ; &6!

r:_a3"!

i

•

•iY

a:":

?•.. ... i:::: -~ f ::

Figure 8. Cached Corrugated Ceramic Vessel (42SA17599) Found in Canyonlands National Park, Utah

Courtesy Midwest Archeological Center, NPS

gathering during the growing season. As several investigators have pointed out, maize might have been eaten green during the growing season (Cordell and Upham 1984; Snow 1991). Green corn consumption could have been integrated into summer hunting and gathering activities-particularly in areas with short growing seasons. Ripened maize would have entailed a more serious commitment to maize production and food storage. Ripened maize served as a critical component of ethnographically documented overwintering strategies in the American Southwest.

CONCLUSIONS

Dynamic aspects of dietary change in the prehistoric American Southwest have usually been explained with respect to the collection or production of plant resources. Until recently, little significance had been ascribed to the role of hunting in forager, collector, and cultivator societies in this region primarily as a result of archaeologists' overreliance on plant-based Great Basin models. Ethnohistorical accounts of Hopi, Zuni, Pima, and Papago horticultural societies have, no doubt, constrained archaeologists' thinking about the potential variation in past feeding strategies. As a result, archaeologists have not, until quite recently, given much thought to high-elevation environments and their role in prehistoric land use in the American Southwest.




This paper has presented a first approximation of an ecological model for ungulates and their human predators in the Colorado Plateau and adjacent mountainous areas of New Mexico and Arizona. Special emphasis has been given to the determinant effects of winter precipitation-specifically snowfall- on resident upland herds of mule deer, elk, bighorn sheep, and, in some cases, bison and antelope. Wildlife ecology studies have identified a number of very powerful determinant relationships that link ungulate behavior and demography to winter precipitation. Archaeologists can make very effective use of winter precipitation, then, as a "proxy" variable for modeling the dynamic aspects of ungulate distribution, abundance, condition, and viability. We will have to make use of these models in order to explain a number of evolutionary developments associated with hunter-gatherers and cultivators in the American Southwest, North America, and the world. We will, no doubt, have to apply these models of ungulate ecology in our future studies of the evolution of agriculture.

The wildlife ecology studies described here are not simply heuristic devices or analog models. They are essential components of a more inclusive explanatory framework. This framework serves to integrate higher-level optimality theory with lower-level physiological, biogeographical, and biometeorological theory. Such a general theory also generates hypotheses and assists in op- erationalizing critical variables. Recent applications of evolutionary ecology in archaeology and anthropology, particularly those that have made use of optimality models, have rarely provided us with insights about human behavior that can be operationalized for empirical testing based on archaeological observations. This limitation has left a number of archaeologists in doubt about the utility of evolutionary ecology in general and optimization models in par- ticular. This paper is meant to redirect our thinking about the role of general ecological theory in archaeology and how it can be used more effectively to tell us not only where to look for relevant empirical evidence but also what such empirical evidence might look like.

NOTE

1. I would like to express my appreciation to Lewis R. Binford (Department of Anthropology, Southern Methodist University); Cal Calabrese, Ralph J. Hartley, Ann Wolley, and Doug Scott (Midwest Archeological Center, National Park Service); Ray- mond Hames, Robert K. Hitchcock, and Beth R. Ritter (Department of Anthropology, University of Nebraska); and June-el Piper and Ron Kneebone (Department of An- thropology, University of New Mexico) for their encouragement, support, and editorial assistance during the development of this paper. James N. Hill (Department of An- thropology, University of California at Los Angeles) provided critical, motivational comments on an earlier version of this paper. I would like to dedicate this article to my parents Mary Ann and Stan Osborn.

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Snowblind in the Desert Southwest: Moisture Islands, Ungulate Ecology, and Alternative Prehistoric Overwintering Strategies