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Clovis Hunting Strategies, or How to Make out on Plentiful Resources

Nicole M. Waguespack; Todd A. Surovell
American Antiquity, Vol. 68, No. 2. (Apr., 2003), pp. 333-352.
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REPORTS

CLOVIS HUNTING STRATEGIES,

OR HOW TO MAKE OUT ON PLENTIFUL RESOURCES

Nicole M. Waguespack and Todd A. Surovell

Traditionally, hunter-gatherers of the Clovis period have been characterized as specialized hunters of large terrestrial manzmals. Recent critiques have attempted to upend this position both empirically and theoretically, alternatively favoring a more
generalized foraging economy. In this paper, the distinction between subsistence specialists and generalists is framed in terms
offorager selectivify with regards to huntedprey, following a behavioral ecological framework. Faunal data are compiled from
33 Clovis sites and used to test the two alternative diet-breadth hypotheses. The data support the older "Clovis as specialist"
model, although some use of s m l l game is apparent. Furthermore, data from modern hunter-gatherers are marshaled to support the theoretical plausibility of specialized large-mammal hunting across North America during the Late Pleistocene.
Tradicionalmente, 10s cazudores-recolectores del periodo Clovis han sido caracterizados como cazadores especializudos en la
fauna mayor. Sin embargo, criticas recientes han tratado de cambiar estaposicidn por una que favorece la economia de recoleccidn generalizada. Este articulo discute la distincidn entre la subsistencia generalizada y la especializuda en te'rminos de la selectividad del recolector respecto a las presas de caza, desde un punto de vista ecoldgico conductual. Se utilizan datos faunisticos
compilados de 33 sitios Clovis para investigar las dos hipdtesis alternas sobre la variedud diete'tica.Los datos apoyan el modelo
tradicional de "especialistas Clovis" aunque aparentemente estos cazadores utilizaron alguna fauna menor. Ademds, datosprovenientes de cazadores-recolectores modernos refiterzan la veracidad tedrica de la caceria defauna mayor a travks de Norte America durante el Pleistoceno Tardfo.

F"

om the initial finds of Clovis points in association with Late Pleistocenemegafauna in the
1930s, Clovis hunter-gatherers have become
synonymous with big-game hunting. In fact, the
lifestyle of Clovis peoples has frequently, and often
solely, been characterized by their unique propensity
to prey upon extremely large-bodiedanimals.As they
hunted not only big game, but Pleistocene rnarnrnothsized game, the "Clovis as hunter" concept has established itself, depending on one's perspective,as either
a proven archaeological fact or as a pervasive and fallacious stereotype. The extent to which Clovis peoples relied upon large game is not just an issue of
subsistence. Their alleged economic emphasison the
hunting of Pleistocenemegafaunahas played an integral role in interpreting their success as colonizers
(Kelly and Todd 1988; Surovell2000), their technological strategy (Frison 1991; Pearson 2001), their
mobility regime (Kelly 1996, 1999; Kelly and Todd
1988), the formation of their archaeological sites
(Meltzer 1993),and ultimately, the extinction of their

.

prey (Alroy 2001; Haynes 2002; Martin 1984;Mosimann and Martin 1975).
Revising the long-standing perspective of Clovis
peoples by characterizing them as generalized foragers would mark a radical departure from the last
65 years of research regarding Clovis lifeways. Some
would applaud this change as a long-overdue revision of an inherently flawed concept (Bryan 1991;
Dillehay 2000: 15-1 8; Gero 1995; Meltzer 1989,
1993;Meltzer and Smith 1986). Others might claim
it a premature and hasty rejection of an established
model (e.g., Haynes 2002). Thls paper evaluates the
theoretical and archaeological implications of the
emergent "Clovis as generalist" and the traditional
"Clovis as specialist" model based on published
archaeofaunalassemblages and on comparisons with
a sample of ethnographic hunter-gatherers.
Clovis as Big-Game Hunters
In the early- to mid-1900s, the classification of Clovis peoples as specialized hunters was initially

Nicole M. Waguespack and Todd A. Surovell Department of Anthropology, University of Arizona, Tucson AZ 85721
American Antiquity, 68(2), 2003, pp. 333-352

Copyright@ 2003 by the Society for American Archaeology

The extensive use of exotic lithic raw materials and an emphasis on biface production (Goodyear 1989. lingering doubts regarding the cultural associationwith numerous megafauna deposits render the quantification and comparison of Clovis kill site faunal assemblages difficult. Figgins 1933. Lundelius 1972). First. mastodon. has led many researchers to doubt the consistency with which Clovis technology is universally found in association with megafauna. deposits throughout North America and Mexico containing any lithic tools in association with Pleistocene megafaunal remains have often. 68. Actualistic studies [Vol. Leonhardy 1966.Haury et al. and/or bison (e.1991. and Lamb Spring (Stanford 1981).334 AMERICAN ANTIQUITY derived from the discovery of numerous sites across North America containing mammoth remains in association with Clovis projectile points (Cotter 1937.g. Once this co-occurrence was firmly established. been attributed to Clovis hunting activities. although the exact timing of extinction of numerous taxa remains questionable (Meltzer and Mead 1985). Johnson 1987. Meltzer 1988. Deuwall-Nuberry (Steele and Carlson 1989).Haury 1953. usually attributed to the greater archaeological visibility of large faunal remains: Most sites were initially paleontological discoveries-the bones of megafauna are visible more readily than artifacts-and the attention to the bone-bearing deposits led to the artifacts and.2001. such as Boaz (Palmer and Stoltman 1975).The majority of critics acknowledge that Clovis peoples killed large game at least occasionally. The frequently questionable relationship between artifacts and faunal remains at many reported proboscidean localities. with an overwhelming bias toward the discovery of largegame kill sites.Although only a handful of these taxa have been recovered from Clovis sites in clear association with artifacts (Grayson 1984). thus to a Paleoindian record largely comprised of kill or scavenging sites [Meltzer 1989:477].Martin 1984. Unfortunately. Sellards 1952). Studies of elephant populations in Africa have led to some troubling observations concerning archaeological associations between proboscideans and humans (Haynes 1988. It is argued that the archaeological record of Clovis suffers from unsystematic sampling. Howard 1935. This argument implies that if more campsites. 1959. In addition to the faunal record. McLean (Ray 1942). or at least "nonkill" sites are discovered.. Haynes 1993. the fact that Clovis projectile points are not consistently found in archaeological deposits associated only with medium-to-small prey has convinced many archaeologiststhat Clovis peoples were big-game specialists. No. The most common points of contention between the two positions are the presumed ubiquity of Clovis pointlmegafauna associations and the viability of big-game hunting as a reliable subsistence strategy for a colonizing population in Pleistocene North America. . but not always. the defining characteristic of Clovis lithic technology is the manufacture of large bifacial projectile points.they would likely present evidence attesting to the consistent utilization of a diversity of faunal and floral species (Gero 1995. 2.Mosimann and Martin 1975). Hudecek-Cuffe 1998). camel. 1993. some researchers accept this temporal coincidence as evidence that the Pleistocene megafaunal extinction was due to overharvesting by Clovis populations (Alroy 1998. However.but argue that the bulk of their diet was comprised of small game and plant resources. Frison and Todd 1986. Haury 1953. A related issue is the reliability of mamrnoth/Clovis associations. Second. 1995). Meltzer and Smith 1986). to theoretical concerns as to whether big-game hunting could have been an economically feasible strategy for Clovis peoples (Meltzer 1993. 1988. Kelly and Todd 1988)have been interpreted as evidence of Clovis peoples' high mobility. Some authors note little or no evidence for largegame hunting in regions such as the Great Basin (Heizer and Baumhoff 1970)or eastern North America (Meltzer 1988). the interpretation of Clovis peoples as big-game hunters has been inferred from ancillary evidence. Clovis as Generalists Critiques of Clovis people as big-game hunters have ranged from accusations that Clovis hunting behaviors provide a medium for perpetuating androcentric bias (Gero 1995. 20031 have established that these weapons are capable of woundingrkilling large prey such as modern elephants (Frison 1989). Despite the many ambiguous associations. the age of Clovis is roughly coincident with the extinction of over 35 genera of large mammals. it must be acknowledgedthat nearly all generally accepted Clovis sites containing well-preserved faunal remains include in their assemblages large-bodied species such as mammoth. a strategy congruent with the pursuit of large game. Laub et al.1995). Johnson 1977.

1987). The diverse environments present during Clovis times and occupied by Clovis peoples would have favored a generalized resource acquisition strategy (Bryan 1991. There is evidence from a number of sites that Clovis hunter-gatherers did include a diverse set of resources in their diet. 1987. Big-game hunting specialization is a sustainable subsistence strategy only when the targeted prey is largelabundant and has high renewal rates (Cleland 1976.Meltzer and Smith 1986). This position runs contrary to the predicted use of resources under optimal foraging models. 1993. small mammals. Big-game hunting specialization is ethnographically rare and can only be expected to occur among foraging peoples in restricted. homogeneous. tortoise. and snake (Haynes and Haury 1982). rather than specialized selection of healthy animals.juveniles and adult kills are represented (Haynes 1991. Hill et al. Formal diet-breadth models (Stephens and Krebs 1986) would predict. Meltzer and Smith 1986). Meltzer 1993. the killing of animals severely weakened by drought or illness. Graham and Kay 1988. large-game specialization is expected to occur only in environmental contexts without other foraging options. the following statements have been used to suggest that big-game specialization is not only unlikely. that large-body-sized animals would be extremely high-ranked prey in any environmental context in which they are available (Hawkes et al. Kelly 1996. 1993). fish. birds. 1992). McNett et al.g. Johnson 1987. For example. Exclusively theoretical arguments against Clovis 335 as big-game specialists are based primarily on ecological models and ethnographically known huntergatherer subsistencebehaviors. Dixon 1999. Three general points regarding the likelihood and potential success of a specialized hunting strategy during Clovis times are consistently cited as reasons against a big-game hunting focus. in addition to the large game found at the Lehner site. 1980. assuming that large game could successfully be captured and reliably fulfill resource needs.Although critics of the Clovis-as-specialistsmodel are quick to point out that the presence of megafauna does not necessarily imply selective hunting (Bryan 199 1. Meltzer 1988.Large-body-sizedanimals are consistently rare compared to the population densities of smaller co-resident species in any given envi- .. African elephants tend to die near sources of water. Dixon 1999:247-250. The few Clovis kill sites with age and sex data for mammoths show various patterns. Presented as foraging constraints. particularly during drought years when catastrophic die-offs can occur. The most straightforward critiques point to the fact that few historically known and extant foraging groups prey exclusively upon big game outside of arctic environments (Johnson 1977. observations of weak and dying elephants during drought years have led to the conclusion that Clovis elephant hunting activities may have been "moribund scavenging" (Fisher 1986). rendering them a poor choice for predation (Meltzer 1993). The Aubrey site also produced a range of small animals. 1991. Haynes and Haury 1982. including jackrabbit. age and sex profiles of catastrophic elephant die-offs at water holes can mimic those of mass kills (Haynes 1988. From this perspective. Johnson 1977.Other sites such as Blackwater Draw (Lundelius 1972)and Lubbock Lake (Johnson 1987) contain small-game faunal assemblages.1993. The presence of small game has led some researchers to conclude that "Paleoindian subsistence data indicate an economic system rooted in general foraging" (Dixon 1999:255). Kornfeld 1988. Ferring 1995. low biodiversity environments (Hayden 1981. The argument that specialized big-game hunting would not have provided a sustainable or reliable strategy for Clovis peoples assumes that the large game were rare andlor potentially dangerous to their human hunters.1995). 3. utilizing a variety of animal food from a wide array of vertebrates both large and small" (Johnson 1991:229).and "Opportunistic hunting appears as a broad-spectrum meatrelated subsistence base. as both catastrophic and isolated individual males. Meltzer and Smith 1986). Winterhalder 1986. Jochim 1981. 1977). utilizing plants. Finally. the assemblage contained various small game. Lee 1968. including abundant evidence for the exploitation of tortoises (Ferring 1995). As many Clovis sites are located in or near springs. Meltzer 1988. and reptiles (e.REPORTS Notably. females.1989. Johnson 1991. Meltzer 1988. but theoretically impossible: 1. 2. Saunders 1977. draws and playas. the potential for fortuitous associations between mammoths and artifacts is high. Also. 1982. Dillehay 2000:28-34. Meltzer 1993). it seems reasonable to also question whether the presence of small game alone provides an adequate indicator of a generalized foraging strategy.

too inflexible to cope with the environmental diversity encountered. Meltzer and Smith (1986) have argued that in such circumstances. is the relative degree to which these taxa are exploited when encountered. 20031 of the features of a new region because that other group did not exist? Entering an unpopulated continent. They suggest that large-game hunting is successful only in specific ecological niches and that generalized foraging is inherently more "adaptable" to a diverse range of environments: Specialists are more efficient and more productive on their own ground and are thus in some sense "stable" so long as the rules of the game are unchanged. [Meltzer and Smith 1986:8]. Generalized vs. however. an understanding of animal behavior. generalists should utilize a broad range of species.a generalizedforaging strategy would be favored over specialized hunting. Clovis big-game hunting is then construed as a nonviable subsistencestrategy for a colonizing population. What means of adapting to local resource stress would be available if Paleoindians could not have depended on another group's knowledge [Vol. which they suggest would attest to the frequent exploitation of small game.Contrary to this position. this does not appear to have deterred modem huntergatherers from pursuing them (Duffy 1984. what is most important from a human subsistence perspective is the ratio of large-body-sized animals to hunters/consumers. a forager will target for predation. the degree to which certain taxa are not exploited upon discovery. Another possibility is that Pleistocene megafauna such as mammoths and mastodons posed such a threat to human life that they would be hunted only when chance favored humans. Stephens and Krebs 1986). while specialists should exploit a more narrow range. Kelly and Todd (1988) argue that large-game hunting is one way foragers can successfully cross-cut environments.. but it is an empirical issue if large-game populations were at densities too low to be consistently preyed upon. On the whole. More important. They argue that a generalized "take what you can" subsistence approach facilitated the rapid expansion of migrating Clovispopulations into new landscapes. As used here they refer to ends of an idealized continuum and concern decisions regarding which prey. Without reliable estimates of Pleistocene megafauna and Clovis population densities. . In this light. among those available. and (3) theoretical concerns regarding the plausibility of a specializedhunting strategy. the most critical variable is selectivity. According to the optimal diet-breadth model.336 AMERICAN ANTIQUITY ronment (Colinvaux 1978. generalists as defined here tend to use. a broad range of taxa when encountered. can be generalized and accommodated to new territories [Kelly and Todd 1988:234]. or at least attempt to utilize. . . a forager can maximize return rates by focusing on taxa whose return rates exceed the average environmental return rate (Charnov 1976. Ultimately. Marks 1976). Peters 1983). (2) negative evidence.e. Specialized Hunting To adequately address the issue of subsistence specialization requires defining the terms "specialist" and "generalist.Taking low-ranked taxa only serves to lower . Climatic and biotic changes give specialized systems a long-term evolutionary liability . "unmapped tracts of land. although compelling. critics argue that large game were too infrequently encountered by Clovis peoples to provide a reliable nutritional source. . can only be resolved if the many "missing" Clovis sites with abundant small game are found. though by no means perfectly transferable. The faunal assemblages of known Clovis sites and the conditions that permit specialized hunting strategies can be addressed with the currently available evidence. 68. Although there are ethnographically documented examples of hunters being wounded (occasionallymortally) by elephants. opponents of the Clovis-as-specializedbig-game-hunter interpretation base their arguments on (1) the presence of small game in archaeological assemblages.Appeals to negative evidence. However. A lifeway suitable for this task is one that placed primary reliance on faunal rather than plant resources. Certainly smaller mammals would have been more abundant in environments occupied by Clovis peoples.i. In sum. while specialists tend to ignore many of the species they encounter in favor of pursuing a more limited high-ranked suite. . Unlike much information on plant resources. The widespread distribution of Clovis points throughout North America indicates that Clovis peoples occupied an environmentally diverse territory. 2. that is not their intended meaning in this discussion. early Paleoindians needed a system which allowed them to utilize unoccupied. No." Although the terms often imply two pure strategies.

and low-rankedtaxa are used is the critical distinction between specialized and generalized strategies. overall return rates. For specialists. it follows that the number of taxa utilized is not in itself a good measure of subsistence specialization. mobility patterns. if environmental and species-specific return rates are allowed to vary. as in a risk-sensitive model (Winterhalder et al. or where little variability exists in return rates among prey items. Predicted faunal assemblages for specialist and generalist hunter-gatherers.-Also. the relative degree to which high. an unproductive activity from an evolutionary standpoint. if high-ranked items are temporally scarce the average environmental return rate effectively drops. a proxy measure must be used. there would be ample reason to expect that even hunter-gatherers that tend toward a specialist strategy would occasionally take low-ranked prey. there should not be a positive correlationbetween encounter rates (as estimated by prey population densities) and archaeological abundance.REPORTS 337 Encounter Rate 0 0 0 Large Prey Medium Prey Small Prey Figure 1. large game would dominate their faunal assemblages. Since generalists will tend to exploit most prey items they encounter. Therefore. prey frequencies in the archaeological record shouldbe positively correlatedwith encounter rates (Figure 1). however. and high-ranked taxa are encountered frequently. A generalized strategy would be expected in environmentswhere high-ranked taxa are infrequently encountered.Since encounter rates for prehistoric prey taxa are unknown. and access to hunting technology. According to the dietbreadth model. Because encounter rate is primarily a function of prey population density. Finally. high-ranked. However. The classic diet-breadth model predicts that a species will either always be taken or always be ignored.This does . Returning to the issue of selectivity. Consequently. but relatively rare. Due to these situational and contextual contingencies. 1999). there would likely be a negative correlation since high-ranked prey are generally large-bodied animals that also tend to have low population densities and therefore low encounter rates. skill. where high-ranked taxa arerarely successfully captured. specialized subsistence strategies should be present in environmentswhere return rates for highly ranked prey species far exceed those of low-ranked items. clear predictions can be made for the archaeological record produced by specialists and generalists. if Clovis foragers behaved as specialists.estimated population densities can provide a reasonable approximation. In fact. For example. and it will likely be exploited. the "zero-one rule" (Stephens and Krebs 1986:20-21). optimal diet breadth should vary for differentsegments of a foragingpop- ulation with respect to age. if a forager encounters a low-ranked prey item in an unusually favorable circumstance with minimal handling costs. the effective return rate for that animal is enhanced. bringing low-rankedprey types into the diet (Haynes 2002).

Although precise quantitative estimates of the population densities of late Pleistocene mammals are not possible. and grassland species. due to inevitable energy losses that occur at higher trophic levels (Colinvaux 1978). and evenness. To this end. Figure 2). our MNIs are based on the number of species present within a given taxonomic grouping. For simplicity. 2001). which have effectively documented temporal trends in the diet breadth of prehistoric foragers (Broughton and Grayson 1993. and Eastern Woodlands). turtles1 . lack fluted points. The data demonstrate a clear relationship between body size and population density (Table 1. in relation to Clovis.g. Although some have argued that the record is unduly biased by an overrepresentation of megafauna kill sites. homeotherms)..g. and this general relationship should exist independent of environment (e. it is predicted that either a negative correlation or no correlation will exist between archaeological species abundance and natural abundance based on body size. assuming that a highly selective subsistence strategy favoring the use of large terrestrial game was the basis of Clovis subsistence economy. but show evidence of human interaction with extinct fauna. 2. which range in body size from 30 g (Common Dormouse) to 5. Grayson et al. the faunal assemblages of 33 archaeological sites were tabulated by species presence1 absence and when possible minimum number of individuals (MNI) (Figure 3. while only presence is noted for others. If Clovis hunter-gatherers were generalists. species diversity. necessarily predict that small game would not have been utilized. desert. herbivores). Grayson and Delpech 1998. Foraging encounter rates can then be expected to vary consistently with regard to prey size based on population densities. Great Plains. For example. Thus. mammothlmastodon. it is reasonable to assume that body [Vol. carnivores exist at lower population densities than herbivores of comparablebody size. the archaeological abundance of prey taxa should be positively correlated with natural abundance. 20031 size can serve as an inverse proxy measure of population density and therefore encounter rates. Faunal data for the remaining sites were recorded simply as presentlabsent for each taxon. existing at the same trophic level (e. it is abundantly clear that proboscideans would be expected to exist at far lower densities than small-game species. lagomorphs. and even the data from some of these sites are problematic. such as rabbits androdents. the archaeological record is the only direct source of information regarding Clovis hunting behaviors. Estimating Clovis Diet Breadth As discussed above. equal amounts of biomass can be consumed by many individuals of a small body size or fewer large-bodied individuals. 6th edition (Nowak 1999). the methodology used here is designed to evaluate the selectivity expressed in prey choice decisions among roughly contemporaneous foragers and their faunal assemblages. Relating prey selectivity to the natural abundance of available prey species creates a methodological framework for examining diet breadth zooarchaeologically.. or in a few cases..338 AMERICAN ANTIQUITY not. Sites were included in the sample if they contained Clovis diagnostics. rodents. This correlation results from the simple fact that for animals with similar metabolic processes (e. In these cases. Unlike measures that rely primarily on the relationshipbetween assemblage size. the more specialized foragers are. 68. There are many intervening variables that serve to muddle this relationship. it should be equally applicable for the Great Basin. 2001. MNIs are sometimes reported for some taxa. Table 2). For example.. reliable MNI values for all taxa present could only be obtained for 15 site assemblages. only that these resources would be hunted less frequently than expected in relation to encounter rate.g. The sample includes forest.g. Due to inconsistencies in the literature with regards to the quantification of fauna. A global sample of body size and population density data for 36 primarily herbivorous mammalian taxa was taken from Walker'sMammals of the World. Estimating Prey Population Densities Although seemingly we have substituted one unknown (encounter rate) for another (prehistoric prey population densities). Peters (1983:164-183) reports similar relationships for a broader array of taxa. the more likely their prey assemblages will deviate from the natural abundance of each species hunted. No. For example.000 kg (African Elephant). it is argued here that it is possible to roughly estimate relative animal population densities for any ecosystem in the past using basic ecological principles and global relationships between prey body size and population density. however. certain species were aggregated into general taxonomic groupings (e.

15 1. Density Page # 1.000 963.0 93. (Nowak 1999).3 115.339 REPORTS Table 1.3 preference was given to the recording of large.25 4. Mass Scientific Name Common Name Order Rodentia Common Dormouse Genus Abrothrix African Bush Squirrels Northern Pygmy Gerbils Coney Rat Red Squirrels Plains Visacha North American Porcupine Capybara Genus Muscardinus Genus Abrothrix Genus Paraxerus Genus Gerbillus Genus Reithrodon Genus Tamiasciurus Lagosturnus rnaximus Erethizon dorsatum Hydrochaeris hydrochaeris Order Lagomorphu Cottontail Rabbits Bushman Rabbit Jack Rabbits (Hares) Genus Sylvilagus Bunolagus monticularis Genus Lepus Order Artiodactyla Dik-diks Water Chevrotain Javelina Gazelles Vicuiia Impala Pronghorn Reedbucks Blue Sheep Axis Deer Ibex White-Tailed Deer Wart Hogs Gemsbok Hartebeest Wapiti Greater Kudu Moose African Buffalo Genus Madoqua H)lemoschus aquaticus Pecari tajacu Genus Gazella Vicugna vicugna Aepyceros melampus Antilocapra americana Genus Redunca Genus Pseudois Genus Axis Capra Ibex Odocoileus virginianus Genus Phacochoerus Genus Oryx Alcelaphus bilselaphus Cervus elaphus Tragelaphus strepsiceros Alces alces Syncerus caffer Order Perissodactyla Burchell's Zebra Black Rhinoceros Equus burchelli Diceros bicornis 1726-1729 1722-1723 1733-1738 Order Proboscidea 994-998 Indian Elephant Elephas muximus 998-1004 African Elephant Loxodonta africana Note: Data from Walker's Mammals of the World. tortoises. amphibians. To maximize the number of sites and by using an extremely lenient measure of cultural association-simple presence in the assemblage-the quantity of all taxa in each assemblage was tabulated. no Pop.18 4.. The presencetabsence of taxa across all 33 sites . 6th ed. the sample is probably preferentially skewed in favor of small game.3 0. and carnivores. and insectivores). Animal Body Mass and Population Density. In fact. Although the inclusion of both the presence and MNI data of species with undoubtedly questionable cultural association skews the distribution of taxa.g. see Table 2 for all groupings). rodents.060 5.6 1.or small-body-sized animals. by counting the presence of numerous small species that are less likely to have been utilized as food sources (e.

(14) Gault. (19) Hiscock. (25) Guest. Nam. 68. (18) Holcombe Beach. 2. Regression line: log. (24) Wacissa River.340 AMERICAN ANTIQUITY [Vol. (5) Charlie Lake Cave. (1) Mannis. (7) McLean. . (10) Miami.o (Population Density) = 1. (11) Domebo. (6) Blackwater Draw. Animal population density versus body mass. (26) Little Salt Spring.962 + 4 3 6 x log. (15) Kimmswick. Leikem. Escapule. (16) Boaz. Map of sites included in this study. (12) Lange Ferguson. (17) Schaefer. (Body Mass) (data from Nowak 1999).20031 Body Mass (kg) Figure 2. (2) Murray Springs. (23) Whipple. Lewisville. (9) Lubbock Lake. (20) Martins Creek. Hebior. Lehner.. (13) Aubrey. Figure 3. (4) Dent. Both axes are log-scaled. (3) Colby.(21) ShawneeMiinisink. (22) Bull Brook. (8) Kincaid. No.

REPORTS " 8C x= e. d eg - 10 B + 6' -5 .r. k .m . NG. ' 9 2 1 Amphibian Bird \ Fish \ Alligator Snake TurtleiTortoise > Insectivore Other Rodent > Muskrat > Armadillo Lagomorphs Other Carnivore Bear Glyptodont Sloth Other Ungulate Peccary Tapir Camelid Equid Bison Proboscidean \'O - P- 2 - .- v.\ N - m \ \ N 2 - N N . P A + 3 - E V .

Alongside typical mammalian faunas. turtleltortoise and birds (30 percent). birds. followed by other ungulates (45 percent). respectively. and glyptodonts (Size Class 3). two general conclusions can be drawn. Rare speciesinclude sloth. The relatively high frequency of rodents and carnivores is likely artificial since little clear evidence of their use as a food source has been documented for Clovis peoples. and Rodents and Insectivores (Size Class 5) (see Table 3). First.tapir. Table 3). Either mammoWmastodon or bison are found in 88 percent of assemblages. Lagomorphs and Armadillos (Size Class 4). and 42 percent of sites contain both. muskrats. equids.9. sloths. Mammoths and mastodon are most common. glyptodont. and lagomorphs (27 percent). . when small-game species are present in an assemblage.037) providing strong support for the Clovis-as-large-game-specialist model. To standardize for varying numbers of taxa in each category. which they are in most but not all cases. (Figure 4) shows that mammoth and mastodon are the most frequently occurring species.3 individuals each. other rodents. the largest species occur in the greatest numbers (Figure 6a. the following categories ranked from largest to smallest were created: Proboscideans (Size Class l). On average. and camelids. fish. The relationship between body size class and MNI value shows a strong. and alligator. In order to analyze the data with regard to prey specialization and diet breadth. and reptiles) and carnivores.Interestingly. the presence of a species was assigned an arbitrary MNI value of one. This is followed by an average of 12. the most consistent component of Clovis faunal assemblages is the presence of mammoth/mastodon (Figure4). two-tailed significance. and equids. amphibians. Second. 2. categories of taxa were further aggregated by general body-size classes. Other ungulates. and camelids. represented by at least 9 1individuals. Bison/ EquidICamelid (Size Class 2). a broad diversity of species is present (Table 2. Other ungulates.20031 V Figure 4. and reptiles are not uncommon constituents of Clovis sites. the largest size group. p = . In order to include the presencelabsence data for analysis. birds.3 individuals. Three species of proboscideans. Figure 4). equids. Both the number of sites in which speciesof each of these five categories are represented and total MNI values were tabulated. Least abundant are the remaining size classes of rodents and insectivores with 3. "other rodents" (MNI = 77) and turtles and tortoises (MNI = 67) are the next most common taxa.1 individuals per species. Excluding all non-mammalian species (birds. the total MNI for each taxonomic grouping was divided by the total number of species included in each size class (Table 3). Bison occur in 52 percent of bone-bearing sites. fish. sloths. and glyptodonts are represented by 4. 68.1 individuals per species. followed by bison. Based on presencelabsence data alone. Percentage of Clovis sites (n = 33) containing each taxonomic grouping. These findings are mimicked by MNI data (Figure 5). No.AMERICAN ANTIQUITY [Vol. Four species of lagomorphs and armadillos are represented by an average of 6. present in 79 percent of all assemblages.2 individuals for 9 species of bison. statistically significant negative correlation (Spearman S p = -. are represented by an average of 30.

assuming a generalist "take what you encounter" strategy (Table 3). In our sample. Stiner et al. Proboscideans appear in 22. Since this method only takes into account numbers of species and not numbers of individuals in each taxonomic grouping. Both taxa are characterized by low reproductive rates resulting from delayed reproductive maturation (Haynes 1991. and least abundant taxa are the most consistent members of Clovis faunal assemblages. In other archaeological contexts high frequencies of chelonians have been used as evidence of harvesting behaviors of human groups with low population densities (Stiner et al. for which there is clear evidence of Clovis utilization (e. Using a Spearman's rho correlation (p = -1. equids.0. For example. a slightly different approach was taken to avoid problems of autocorrelation. and Lehner). 2000). tortoises. the pres- encelabsence data also provide strong support for the Clovis-as-specialist model since a strong negative correlation between body size and presence in Clovis assemblages is suggested.g. while rodents and insectivores are present in 19 fewer sites than expected. Therefore. the expected value is 3. These are essentially predation avoidance mechanisms since both adult turtles.1 more sites than expected. Total MNI by taxonomic group for 33 Clovis sites. they are likely to be high-ranked resources. sloths. 2000). Lagomorphs and armadillos are present in 4. as groups with more taxa should have more opportunities to enter archaeological sites.3 sites. a minimum of 67 individual turtles or tortoises is represented. Aubrey.1 more sites than expected. One of the nonmammalian small-game categories not examined in relation to natural abundance is turtles and tortoises. bison. If an invasive .. it is biased toward smaller. For each size class. Interestingly. consisting of three species. and the expected values for the remaining taxa were set proportionately. despite relatively small body size. turtles and tortoises do the same by directing energy into the growth of bony shells. and camelids appear in 9. making both chelonians and proboscideans especially susceptible to overharvesting and extinction (Mithen 1993. Lubbock Lake. The expected value for rodents and insectivores was set to 33 sites (the total sample). and the largest. for proboscideans. and glypotodonts are present in 8. Figure 6b).7 more sites than expected. The expected value for each size class was standardized to the most taxonomically abundant group (rodents and insectivores. While elephants postpone reproduction by investing energy early in life into growth to reach a large-body size. and other ungulates. least diverse. p = 0). Since turtles and tortoises are easily captured with low handling costs. Shine and Iverson 1995). this pattern is statistically highly significant.6 more sites than expected. Comparison of the observed and expected number of sites shows that the largest taxa are overrepresented in Clovis sites (Table 3. the expected number of sites in which each group should be present was calculated. To investigate the relationship between body size and taxa presence and absence. and elephants have few natural predators. n = 30 species). turtles and tortoises share many life history traits with proboscideans. or one-tenth the diversity of rodents and insectivores. more abundant species.REPORTS Figure 5.

in areas in which the bulk of primary biomass cannot be consumed by humans but is accessible to herbivores (i.4 9 4.6 Rodents and insectivores 5 30 93 3. their subsistence options are perhaps significantly more limited than the constraints on hunter-gatherers of the past. Meltzer and Smith 1986).3 4.344 [Vol.3 26 Actual n sites 22.9 18 8. 20031 AMERICAN ANTIQUITY Table 3. But these patterns are drawn from modern and historically known hunting and gathering groups.The percent of the diet comprised of plantlsmall terrestrial fauna and larger game resources was tallied. Other ungulates. equids. arctic ecosystems).7 Act. peoples (reviewed in Kelly 1995:66-73)... Since 9. however. Kelly 1995.1 33 14 -1 9 most recent foraging populations occupy primarily "marginal" environments and must coexist with neighboring nonforaging societies. and camelids 2 9 110 12. n sites Note: tSee text for explanation of calculation. For Clovis populations living in late Pleistocene North America. land-use options would have had few limits since population densities would have been extremely low. and less commonly. hunting specialization may have been possible in environments where this strategy does not and could not exist today. 1993. environment. and glyptodonts 3 9 37 4. hunter-gatherers documented in the modern era occupy ecosystems that have been inhabited by humans for thousands of years and they likely exist at relatively high population densities (as compared to Clovis). 2. If big-game hunting specialization is a viable strategy in any environment as long as human population densities are low relative to available game. As previously discussed. In addition. then the prey duo of elephants and tortoises is not unexpected. . To examine the relationship between reliance on hunting.1 species. theoretical arguments launched against Clovis-as-specialists frequently point to the fact that ethnographically known examples of foragers with specialized hunting economies are geographically limited to arctic and grassland environments (Meltzer 1988. In such a situation of relatively few people and abundant big-game availability.3 Bison.e.I 9.Exp.e. is expected to move into a niche with minimal interspecific competition. Murdock 1981).9 19 9. large-game specialization is simply not possible because there are too many people and/or too few large animals. and do not necessarily reflect the range of possible subsistence behaviors of the past. hunter-gatherer groups who derive more than 50 percent of their food resources from hunting generally occupy environments with low mean effective temperatures (i. in this case prehistoric humans in Pleistocene North America. 68. a situation that has existed for perhaps thousands of years. then there is little reason to presume that Clovis peoples could not have been specialists simply because they occupied an ecologically diverse environment. No. Populations of hunter-gatherers that derive all of their subsistence resources from foraging were collected from the Atlas of World Cultures(Murdock 1981).1 Lagomorphs and armadillos 4 4 25 6. Clovis Assemblage Data Summarized by Body Size Size Class N Taxa Total MNI MNI per taxon Proboscideans 1 3 91 30. the distribution of Clovis faunal assemblages appears more congruent with a specialized hunting strategy. in tropical ecosystems where foragers trade meat for plant products with local. In most contemporary environments. and population density. Although potentially the result of biased archaeological visibility and recovery. With few exceptions. grasslands and tundra). a sample of 92 foraging groups was examined. the environment occupied and population density for each group was recorded (data from Kelly 1995: Table 6-4). Also. Is Big-Game Hunting Possible? The Clovis faunal record certainly implies the extensive and selective use of large-bodied prey.2 Expected n sitest 3. The ethnographic record does evidence that a strong relationship exists between the proportion of subsistence resources derived from hunted game and environmental context (Keeley 1988. . nonforaging. Some critics. sloths. claim that specialized hunting was simply not a viable subsistence strategy for Clovis peoples.

assuming a generalist subsistence strategy (two-tailedSpearman's p = .037).397. and rifles would have decreased handling costs. The average dependence on larger game decreases with greater population density for all environments (Spearman's p = -. p = . as noted by Meltzer (1993). Hunters of the Great Plains are an interesting exception because. the only exceptions being the Mbuti and groups of the North American Great Plains who both tend to depend more on hunt- ing than would be predicted by their population densities. p < .9. As predicted by the diet-breadth model. (a) (top) Prey size class versus total MNI per species. . they were armed with rifles and had the use of the horse. the actual minus the expected number of sites per taxonomic grouping. Not surprisingly. the introduction of the horse and gun should have caused a contraction of diet breadth because greater mobility would have increased encounter rates with large mammals.1 . A similar effect was observed with the introduction of the snowmobileto the Cree (Winterhalder 1981). dependence on plants and small fauna increases with population density and is statistically significant (Spearman's p = . notably bison. p < . divided by the total number of species in that group (two-tailedSpearman's p = -.68.001).001). Plots of log population density (people per 100 km2) for the sampled hunter-gatherer groups and their reliance on both plants/small fauna and large game are presented in Figure 7a and 7b. calculated as the sum of MNIs for all taxa in each size-class grouping. (b) (bottom) Preysize class vs.REPORTS Size Class Size Class Figure 6. p = 0).

A problem of equifinality arises when these results are compared with the findings of others who have correlated dietary hunting dependence with environmental parameters (Binford2001. Dividing the sample into groups who derive more than 46 percent of their subsistence by hunting from those who do not (Figure 8).1995. The mean population density value for groups with greater than 46 percent reliance on hunting is . Figure 7.001).001).001).5 people per 100 These findings cannot directly implicate Clovis peoples as big-game hunters. Spearman's p = 397. km2. population density for a global sample of hunter-gatherers (n = 92.p c .AMERICAN ANTIQUITY pol. (b) (bottom) proportion of large game in the diet vs.25 people per 100km2.p < . population density for the same groups (Spearman's p = -. Keeley 1988. 68. a two-tailed t-test establishes significant differences between their mean population densities ( p < . rr o o arr r - x ex X m r r -- -m--- Population Density (persons/100 km 2. . 2. No. but it does establish that hunting specialization can and frequently does occur at low population densities. and for less than 46 percent it is 37.20031 ' TropicaliTcmpcratc Dcscrt TropicaliTcmpcratc Forc st CalifomiaNorthwcst Coast A Plains Population Density (persons/100 km 2.68. (a) (top) Proportion of plants in the diet vs.

they moved into a more generalized subsistence strategy. for example. We argue above with respect to the dietbreadth model that large-game specializationshould result from relatively high encounter rates with large game. 347 We argue that large-game specialization is only seen in recent times in areas where there are few other subsistence options. but provides one important source of evidence for the examination of Clovis prey choice. allowing these animals to attain and maintain high population densities.. Therefore. Although prey densities will ultimately be controlled by environmental productivity. Differential recovery and preservation of animal remains that bias the record toward large animals are possible influenceson the Clovis record that currentlyremain difficult to assess. independent of environmental parameters. environment. should be limited by food supplies. Many potential biases exist in the dataset. Percentage of large game in the diet vs. of population density.MI). Data from Kelly (1995:Table 6-4) and Murdock (1981). we are not willing to assume that this bias is so extensive as to reverse our findings. eventually allowing for higher human population densities. resulting in a sticky problem of establishing causality. The zooarchaeological data collected for the current analysis are admittedly imperfect.a hunting-based subsistence economy seems highly plausible. population density for a worldwide sample of hunter-gatherers (n = 92. two-tailed t-test. while acknowledging the existence of these nonrandom factors influencing the dataset.Furthermore. Nonetheless. hunter-gatherers who depend entirely on hunting will exist at far lower densities than those who adopt a more "vegetarian" lifestyle since hunters only exploit a small fraction of the available edible biomass in an ecosystem. have shaped the modern ethnographic record of large-game hunting. Kelly 1995. whichis ultimately limited by subsistencepractices. regardless of the diverse environments they occupied. human population densities can be modified by adjusting "territory" or range size. The inability to switch subsistence strategiesin the arctic. and subsistence). humans chose to be large-game specialists early on. subsistence should ultimately be a product of the density of prey coupled with the density of human hunters. These factors. for the purposes of this study. but due to increasing numbers of people and decreasing numbers of prey. We have framed the contrast between hunting specialization and generalization on the degree of selectivityexpressed in prey choice in amanner consistent with optimal foraging theory. we suggest. In other biomes where large animals were readily available.p < . Conclusions Although there has been much debateregarding Clovis diet breadth and subsistence strategy. Meltzer 1993). Absolute numbers of humans. three independent variables are collinear (population density. An argument could be made that environment is the proximate cause of subsistence practices and population density. as long as Clovis peoples maintained low population densities and large game was available and regularly encountered. a vast majority of the primary biomass present is accessible to large ungulates (due to the paucity of woody vegetation). that of top carnivore. has resulted in one of the very few areas where humans exist at extremely low population densities because they were forced into avery narrow subsistenceniche. few attempts have been made to quantitatively analyze Clovis faunal assemblages. Note Y-axisis the log.REPORTS Percentage of Large Game in Diet Figure 8. although the environments where large-game specializationoccurs in recent times are characterized by relatively low primary production. In other words. however. That is. Therefore. Selectivity has been defined as the relative degree to which certain .

rather than generalized. Adaptive Change. Specialists tend to bypass numerous potential prey in favor of higher-ranked resources. early Paleoindian population densities were undoubtedly low. Brush. N. and published. 2. Clovis huntergatherers often ignored opportunities to harvest smaller game species. Current Research in the Pleistocene 11:1618. Grayson 1993 Diet Breadth. Fisher 1993 Research on the Dent Mammoth Site. This does not mean that Clovis existed by mammoth alone. 1998 Putting North America's End-Pleistocene Megafaunal Extinctions in Context:Large-Scale Analyses of Spatial Patterns. human population density is a critical variable.348 AMERICAN ANTIQUITY taxa are not exploited upon encounter. This conclusion is drawn from the relative abundances of a range of taxa represented in Clovis sites. and F. J. Vance Haynes.. Binford. the taxa most frequently occurring in sites and occurring in the greatest numbers are those that would have been infrequently encountered by Clovis huntergatherers. However. and other large animals. MacPhee. and one anonymous reviewer for their helpful comments and insightful suggestions regarding this work. Low population densities coupled with frequent residential mobility over large distances provide a context where large-game specialization is not only possible but also likely. Robert Kelly. 2001 A Multispecies Overkill Simulation of the End-Pleistocene Megafaunal Mass Extinction. Following Kelly and Todd (1988). however. Brush. The quintessential generalist. and therefore have relatively high handling costs for a relatively small payoff. Unlike the expectations for a generalist strategy. 2001 Constructing Frames of Reference: An Analytical Methodjor Archaeological Theoly Building Using HunterGatherer and Environmental Data Sets. and to Jeffrey Saunders who generously allowed the use of his unpublished faunal inventories of the San Pedro Clovis sites. Since Clovis represents the colonization of an uninhabited landscape or slightly postdates that event. the Clovis faunal record points to a clear conclusion-large-game specialization. we value their patience and contribution to our own thoughts regarding Clovis subsistence. Extinction Rates. 68. are not so easily taken since they have evolved sophisticated defense mechanisms to elude numerous predators. until these sites are discovered. C.. likely in favor of obtaining a higher-ranked resource. H. Current Research in the Pleistocene 10:63-65. we are very much indebted. and the White Mountain Faunas. M. birds. We would like to thank Mary Stiner. on the other hand. L. based on estimated encounter rates. R. subsistence emphasizing large-game hunting is not restricted to any particular environment (except those lacking large game). Mary Lou Larson. Kluwer Academic/Plenum Publishers. or at least regularly encountered (to provide sufficient caloric yields). excavated. it should come as no surprise that these taxa seem to have only played a minor role in Clovis subsistence. Gratitude is also extended to Julio Betancourt and Maria Nieves Zedeiio who translated the abstract. would exploit all or nearly all prey encountered. References Cited Alroy. Smith 1994 The Martins Creek Mastodon: A Paleoindian Butchery . Based on the analysis of 33 faunal assemblages it has been argued that Clovis hunting behaviors appear more closely aligned with a specialized.Despite their presentation here as alternative strategies. and Size Distributions. and F. pp. Berkeley. Jr. N. K. 105-143. and D. J. Acknowledgments.. and has always encouraged us to pursue our interests. Journal ofArchaeologica1Science 20:33 1-336. University of California Press. and is perhaps the ultimate strategy for maneuvering among diverse environments. 20031 erers show a clear increase in hunting of large animals as human population densities decline. Broughton. fish. and other small game taxa.. Nor does it imply that Clovis peoples only hunted mammoth. Although we acknowledge the potential for hundreds of missing Clovis sites with thousands of individualsof small-game species. New York. In Extinctions in Near Time. David Overstreet. Data from modern hunter-gath- [Vol. For her enthusiasm in volunteering and for her companionship. There is clear evidence that they did exploit small mammals and reptiles on occasion. Sincelarge-game specialization is only possible when people are few and large animals are many. Although some of these commenters ultimately disagree with our interpretations. and D. There is not only empirical evidence that Clovis peoples were specializedpredators of large game. Smith 1994 Immunological Analysis of Flint Flakes from the Martins Creek Mastodon Site. Paul Martin. Therefore. Brunswig. Rabbits. strategy. M. C. Newman. edited by R. Science 292:1893-1896. David Andersen. they are intended only to describe two possible extremes of a continuum. bison. Marcel Kornfeld. but this characterization also receives some theoretical support. Turtles and tortoises are one clear example of small game taxa that Clovis foragers may have regularly taken upon encounter since handling costs are minimal for such slow-footed prey. And a special thanks to Cherie Freeman who helped compile data for this study. No. Marvin Kay. R. rodents..

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