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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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Tue Jan 29 21:28:40 2008



Nicole M. Waguespack and Todd A. Surovell

Traditionally, hunter-gatherers of the Clovis period have been characterized as specialized hunters of large terrestrial manz-
mals. 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 sup-
port 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 recolec-
cidn generalizada. Este articulo discute la distincidn entre la subsistencia generalizada y la especializuda en te'rminos de la selec-
tividad 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, datosprove-
nientes de cazadores-recolectores modernos refiterzan la veracidad tedrica de la caceria defauna mayor a travks de Norte Amer-
ica durante el Pleistoceno Tardfo.

om the initial finds of Clovis points in asso- prey (Alroy 2001; Haynes 2002; Martin 1984;Mosi-
ciation with Late Pleistocenemegafauna in the mann and Martin 1975).
1930s, Clovis hunter-gatherers have become Revising the long-standing perspective of Clovis
synonymous with big-game hunting. In fact, the peoples by characterizing them as generalized for-
lifestyle of Clovis peoples has frequently, and often agers would mark a radical departure from the last
solely, been characterized by their unique propensity 65 years of research regarding Clovis lifeways. Some
to prey upon extremely large-bodiedanimals.As they would applaud this change as a long-overdue revi-
hunted not only big game, but Pleistocene rnarnrnoth- sion of an inherently flawed concept (Bryan 1991;
sized game, the "Clovis as hunter" concept has estab- Dillehay 2000: 15-1 8; Gero 1995; Meltzer 1989,
lished itself, depending on one's perspective,as either 1993;Meltzer and Smith 1986). Others might claim
a proven archaeological fact or as a pervasive and fal- it a premature and hasty rejection of an established
lacious stereotype. The extent to which Clovis peo- model (e.g., Haynes 2002). Thls paper evaluates the
ples relied upon large game is not just an issue of theoretical and archaeological implications of the
subsistence. Their alleged economic emphasison the emergent "Clovis as generalist" and the traditional
hunting of Pleistocenemegafaunahas played an inte- "Clovis as specialist" model based on published
gral role in interpreting their success as colonizers archaeofaunalassemblages and on comparisons with
(Kelly and Todd 1988; Surovell2000), their techno- a sample of ethnographic hunter-gatherers.
logical strategy (Frison 1991; Pearson 2001), their
mobility regime (Kelly 1996, 1999; Kelly and Todd Clovis as Big-Game Hunters
1988), the formation of their archaeological sites In the early- to mid-1900s, the classification of Clo-
(Meltzer 1993),and ultimately, the extinction of their vis 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

334 AMERICAN ANTIQUITY [Vol. 68, No. 2, 20031

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

Notably, African elephants tend to die near sources as big-game specialists are based primarily on eco-
of water, particularly during drought years when cat- logical models and ethnographically known hunter-
astrophic die-offs can occur. As many Clovis sites gatherer subsistencebehaviors. Three general points
are located in or near springs, draws and playas, the regarding the likelihood and potential success of a
potential for fortuitous associations between mam- specialized hunting strategy during Clovis times are
moths and artifacts is high. Also, age and sex pro- consistently cited as reasons against a big-game hunt-
files of catastrophic elephant die-offs at water holes ing focus. Presented as foraging constraints, the fol-
can mimic those of mass kills (Haynes 1988,1995). lowing statements have been used to suggest that
The few Clovis kill sites with age and sex data for big-game specialization is not only unlikely, but the-
mammoths show various patterns, as both cata- oretically impossible:
strophic and isolated individual males, females,juve- 1. Big-game hunting specialization is ethnographi-
niles and adult kills are represented (Haynes 1991; cally rare and can only be expected to occur
Saunders 1977, 1980, 1992). Finally, observations among foraging peoples in restricted, homoge-
of weak and dying elephants during drought years neous, low biodiversity environments (Hayden
have led to the conclusion that Clovis elephant hunt- 1981; Kornfeld 1988; Lee 1968; Meltzer 1993;
ing activities may have been "moribund scaveng- Meltzer and Smith 1986).
ing" (Fisher 1986), the killing of animals severely 2. Big-game hunting specialization is a sustainable
weakened by drought or illness, rather than special- subsistence strategy only when the targeted prey
ized selection of healthy animals. is largelabundant and has high renewal rates (Cle-
There is evidence from a number of sites that Clo- land 1976; Jochim 1981; Meltzer 1988, 1993).
vis hunter-gatherers did include a diverse set of 3. The diverse environments present during Clovis
resources in their diet, utilizing plants, small mam- times and occupied by Clovis peoples would have
mals, birds, fish, and reptiles (e.g., Ferring 1995; favored a generalized resource acquisition strat-
Graham and Kay 1988; Haynes and Haury 1982; egy (Bryan 1991; Dixon 1999; Johnson 1977,
Johnson 1987; McNett et al. 1977). For example, in 1991; Meltzer 1993).
addition to the large game found at the Lehner site, The most straightforward critiques point to the
the assemblage contained various small game, fact that few historically known and extant foraging
including jackrabbit, tortoise, and snake (Haynes groups prey exclusively upon big game outside of
and Haury 1982). The Aubrey site also produced a arctic environments (Johnson 1977; Meltzer 1988,
range of small animals, including abundant evidence 1993; Meltzer and Smith 1986). From this perspec-
for the exploitation of tortoises (Ferring 1995).Other tive, large-game specialization is expected to occur
sites such as Blackwater Draw (Lundelius 1972)and only in environmental contexts without other forag-
Lubbock Lake (Johnson 1987) contain small-game ing options. This position runs contrary to the pre-
faunal assemblages. The presence of small game has dicted use of resources under optimal foraging
led some researchers to conclude that "Paleoindian models. Formal diet-breadth models (Stephens and
subsistence data indicate an economic system rooted Krebs 1986) would predict, assuming that large
in general foraging" (Dixon 1999:255),and "Oppor- game could successfully be captured and reliably
tunistic hunting appears as a broad-spectrum meat- fulfill resource needs, that large-body-sized animals
related subsistence base, utilizing a variety of animal would be extremely high-ranked prey in any envi-
food from a wide array of vertebrates both large and ronmental context in which they are available
small" (Johnson 1991:229).Although critics of the (Hawkes et al. 1982; Hill et al. 1987; Kelly 1996;
Clovis-as-specialistsmodel are quick to point out that Winterhalder 1986, 1987).
the presence of megafauna does not necessarily The argument that specialized big-game hunting
imply selective hunting (Bryan 199 1; Dillehay would not have provided a sustainable or reliable
2000:28-34; Dixon 1999:247-250; Johnson 1991; strategy for Clovis peoples assumes that the large
Meltzer 1988,1989,1993;Meltzer and Smith 1986), game were rare andlor potentially dangerous to their
it seems reasonable to also question whether the pres- human hunters, rendering them a poor choice for pre-
ence of small game alone provides an adequate indi- dation (Meltzer 1993).Large-body-sizedanimals are
cator of a generalized foraging strategy. consistently rare compared to the population densi-
Exclusively theoretical arguments against Clovis ties of smaller co-resident species in any given envi-
336 AMERICAN ANTIQUITY [Vol. 68, No. 2, 20031

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

Encounter Rate

Large Medium Small

Prey Prey Prey

Figure 1. Predicted faunal assemblages for specialist and generalist hunter-gatherers.

overall return rates, an unproductive activity from an ulation with respect to age, skill, mobility patterns,
evolutionary standpoint. According to the diet- and access to hunting technology.
breadth model, specialized subsistence strategies Due to these situational and contextual contin-
should be present in environmentswhere return rates gencies, it follows that the number of taxa utilized
for highly ranked prey species far exceed those of is not in itself a good measure of subsistence spe-
low-ranked items, and high-ranked taxa are encoun- cialization. Therefore, the relative degree to which
tered frequently. A generalized strategy would be high- and low-rankedtaxa are used is the critical dis-
expected in environmentswhere high-ranked taxa are tinction between specialized and generalized strate-
infrequently encountered, where high-ranked taxa gies. Returning to the issue of selectivity, clear
arerarely successfully captured, or where little vari- predictions can be made for the archaeological record
ability exists in return rates among prey items. produced by specialists and generalists. Since gen-
The classic diet-breadth model predicts that a eralists will tend to exploit most prey items they
species will either always be taken or always be encounter, prey frequencies in the archaeological
ignored, the "zero-one rule" (Stephens and Krebs record shouldbe positively correlatedwith encounter
1986:20-21). However, if environmental and rates (Figure 1).Since encounter rates for prehistoric
species-specific return rates are allowed to vary, as prey taxa are unknown, a proxy measure must be
in a risk-sensitive model (Winterhalder et al. 1999), used. Because encounter rate is primarily a function
there would be ample reason to expect that even of prey population density,estimated population den-
hunter-gatherers that tend toward a specialist strat- sities can provide a reasonable approximation. For
egy would occasionally take low-ranked prey. For specialists, however, there should not be a positive
example, if a forager encounters a low-ranked prey correlationbetween encounter rates (as estimated by
item in an unusually favorable circumstance with prey population densities) and archaeological abun-
minimal handling costs, the effective return rate for dance. In fact, there would likely be a negative cor-
that animal is enhanced, and it will likely be relation since high-ranked prey are generally
exploited.-Also, if high-ranked items are temporally large-bodied animals that also tend to have low pop-
scarce the average environmental return rate effec- ulation densities and therefore low encounter rates.
tively drops, bringing low-rankedprey types into the Consequently, if Clovis foragers behaved as spe-
diet (Haynes 2002). Finally, optimal diet breadth cialists, high-ranked, but relatively rare, large game
should vary for differentsegments of a foragingpop- would dominate their faunal assemblages.This does
338 AMERICAN ANTIQUITY [Vol. 68, No. 2, 20031

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

Table 1. Animal Body Mass and Population Density.

Mass Pop. Density

Common Name Scientific Name Page #
Order Rodentia
Common Dormouse Genus Muscardinus
Genus Abrothrix Genus Abrothrix
African Bush Squirrels Genus Paraxerus
Northern Pygmy Gerbils Genus Gerbillus
Coney Rat Genus Reithrodon
Red Squirrels Genus Tamiasciurus
Plains Visacha Lagosturnus rnaximus
North American Porcupine Erethizon dorsatum
Capybara Hydrochaeris hydrochaeris

Order Lagomorphu
Cottontail Rabbits
Genus Sylvilagus 1726-1729 1.15 963.3
Bushman Rabbit
Bunolagus monticularis 1722-1723 1.25 115.0
Jack Rabbits (Hares)
Genus Lepus 1733-1738 4.18 93.3

Order Artiodactyla
Genus Madoqua
Water Chevrotain
H)lemoschus aquaticus
Pecari tajacu
Genus Gazella
Vicugna vicugna
Aepyceros melampus
Antilocapra americana
Genus Redunca
Blue Sheep
Genus Pseudois
Axis Deer
Genus Axis
Capra Ibex
White-Tailed Deer
Odocoileus virginianus
Wart Hogs
Genus Phacochoerus
Genus Oryx
Alcelaphus bilselaphus
Cervus elaphus
Greater Kudu
Tragelaphus strepsiceros
Alces alces
African Buffalo
Syncerus caffer

Order Perissodactyla
Burchell's Zebra
Equus burchelli
Black Rhinoceros
Diceros bicornis

Order Proboscidea
Indian Elephant
Elephas muximus 994-998 4,060 0.6
African Elephant
Loxodonta africana 998-1004 5,000 1.3
Note: Data from Walker's Mammals of the World, 6th ed. (Nowak 1999).

tortoises, and carnivores; see Table 2 for all group- preference was given to the recording of large- or
ings). To maximize the number of sites and by using small-body-sized animals. In fact, by counting the
an extremely lenient measure of cultural associa- presence of numerous small species that are less
tion-simple presence in the assemblage-the quan- likely to have been utilized as food sources (e.g.,
tity of all taxa in each assemblage was tabulated. amphibians, rodents, and insectivores), the sample
Although the inclusion of both the presence and MNI is probably preferentially skewed in favor of small
data of species with undoubtedly questionable cul- game.
tural association skews the distribution of taxa, no The presencetabsence of taxa across all 33 sites
340 AMERICAN ANTIQUITY [Vol. 68, No. 2,20031

Body Mass (kg)

Figure 2. Animal population density versus body mass. Both axes are log-scaled. Regression line: log,o (Population Density)
= 1.962 + 4 3 6 x log,, (Body Mass) (data from Nowak 1999).

Figure 3. Map of sites included in this study. (1) Mannis; (2) Murray Springs, Lehner, Nam, Escapule, Leikem; (3) Colby;
(4) Dent; (5) Charlie Lake Cave; (6) Blackwater Draw; (7) McLean; (8) Kincaid; (9) Lubbock Lake; (10) Miami; (11)
Domebo; (12) Lange Ferguson; (13) Aubrey, Lewisville; (14) Gault; (15) Kimmswick; (16) Boaz; (17) Schaefer, Hebior; (18)
Holcombe Beach; (19) Hiscock; (20) Martins Creek;(21) ShawneeMiinisink; (22) Bull Brook; (23) Whipple; (24) Wacissa
River; (25) Guest; (26) Little Salt Spring.

8C x=
e; '
2 1

Bird \

Fish \



TurtleiTortoise >

Other Rodent >

Muskrat >


Other Carnivore




Other Ungulate






Proboscidean \'O - P- 2 - - \ N - m \ \ N 2 N N - m - - - v, d - - 10


V .
eg B

AMERICAN ANTIQUITY [Vol. 68, No. 2,20031

Figure 4. Percentage of Clovis sites (n = 33) containing each taxonomic grouping.

(Figure 4) shows that mammoth and mastodon are amphibians, and reptiles) and carnivores, the fol-
the most frequently occurring species, present in 79 lowing categories ranked from largest to smallest
percent of all assemblages. Bison occur in 52 percent were created: Proboscideans (Size Class l), Bison/
of bone-bearing sites, followed by other ungulates (45 EquidICamelid (Size Class 2), Other ungulates,
percent), other rodents, turtleltortoise and birds (30 sloths, and glyptodonts (Size Class 3), Lagomorphs
percent), and equids, muskrats, and lagomorphs (27 and Armadillos (Size Class 4), and Rodents and
percent). Rare speciesinclude sloth, glyptodont,tapir, Insectivores (Size Class 5) (see Table 3). Both the
and alligator. Either mammoWmastodon or bison are number of sites in which speciesof each of these five
found in 88 percent of assemblages, and 42 percent categories are represented and total MNI values were
of sites contain both. The relatively high frequency tabulated. In order to include the presencelabsence
of rodents and carnivores is likely artificial since lit- data for analysis, the presence of a species was
tle clear evidence of their use as a food source has assigned an arbitrary MNI value of one. To stan-
been documented for Clovis peoples. dardize for varying numbers of taxa in each category,
Based on presencelabsence data alone, two gen- the total MNI for each taxonomic grouping was
eral conclusions can be drawn. First, the most con- divided by the total number of species included in
sistent component of Clovis faunal assemblages is each size class (Table 3).
the presence of mammoth/mastodon (Figure4). Sec- On average, the largest species occur in the great-
ond, when small-game species are present in an est numbers (Figure 6a, Table 3). Three species of
assemblage, which they are in most but not all cases, proboscideans, the largest size group, are repre-
a broad diversity of species is present (Table 2, Fig- sented by an average of 30.3 individuals each. This
ure 4). Alongside typical mammalian faunas, birds, is followed by an average of 12.2 individuals for 9
fish, and reptiles are not uncommon constituents of species of bison, equids, and camelids. Four species
Clovis sites. These findings are mimicked by MNI of lagomorphs and armadillos are represented by an
data (Figure 5). Mammoths and mastodon are most average of 6.3 individuals. Other ungulates, sloths,
common, represented by at least 9 1individuals.Inter- and glyptodonts are represented by 4.1 individuals
estingly, "other rodents" (MNI = 77) and turtles and per species. Least abundant are the remaining size
tortoises (MNI = 67) are the next most common taxa, classes of rodents and insectivores with 3.1 indi-
followed by bison, equids, birds, and camelids, viduals per species. The relationship between body
respectively. size class and MNI value shows a strong, statisti-
In order to analyze the data with regard to prey cally significant negative correlation (Spearman S
specialization and diet breadth, categories of taxa p = -.9; two-tailed significance, p = .037) providing
were further aggregated by general body-size classes. strong support for the Clovis-as-large-game-spe-
Excluding all non-mammalian species (birds, fish, cialist model.

Figure 5. Total MNI by taxonomic group for 33 Clovis sites.

To investigate the relationship between body size encelabsence data also provide strong support for
and taxa presence and absence, a slightly different the Clovis-as-specialist model since a strong nega-
approach was taken to avoid problems of autocor- tive correlation between body size and presence in
relation. For each size class, the expected number Clovis assemblages is suggested, and the largest,
of sites in which each group should be present was least diverse, and least abundant taxa are the most
calculated, assuming a generalist "take what you consistent members of Clovis faunal assemblages.
encounter" strategy (Table 3). The expected value One of the nonmammalian small-game cate-
for each size class was standardized to the most tax- gories not examined in relation to natural abundance
onomically abundant group (rodents and insecti- is turtles and tortoises, for which there is clear evi-
vores, n = 30 species), as groups with more taxa dence of Clovis utilization (e.g., Lubbock Lake,
should have more opportunities to enter archaeo- Aubrey, and Lehner). In our sample, a minimum of
logical sites. The expected value for rodents and 67 individual turtles or tortoises is represented. In
insectivores was set to 33 sites (the total sample), other archaeological contexts high frequencies of
and the expected values for the remaining taxa were chelonians have been used as evidence of harvest-
set proportionately. For example, for proboscideans, ing behaviors of human groups with low population
consisting of three species, or one-tenth the diver- densities (Stiner et al. 2000). Since turtles and tor-
sity of rodents and insectivores, the expected value toises are easily captured with low handling costs,
is 3.3 sites. Since this method only takes into account they are likely to be high-ranked resources, despite
numbers of species and not numbers of individuals relatively small body size. Interestingly, turtles and
in each taxonomic grouping, it is biased toward tortoises share many life history traits with pro-
smaller, more abundant species. Comparison of the boscideans. Both taxa are characterized by low
observed and expected number of sites shows that reproductive rates resulting from delayed repro-
the largest taxa are overrepresented in Clovis sites ductive maturation (Haynes 1991; Shine and Iver-
(Table 3, Figure 6b). Proboscideans appear in 22.7 son 1995), making both chelonians and
more sites than expected, bison, equids, and proboscideans especially susceptible to overhar-
camelids appear in 9.1 more sites than expected, and vesting and extinction (Mithen 1993; Stiner et al.
other ungulates, sloths, and glypotodonts are pre- 2000). While elephants postpone reproduction by
sent in 8.1 more sites than expected. Lagomorphs investing energy early in life into growth to reach a
and armadillos are present in 4.6 more sites than large-body size, turtles and tortoises do the same by
expected, while rodents and insectivores are present directing energy into the growth of bony shells.
in 19 fewer sites than expected. Using a Spearman's These are essentially predation avoidance mecha-
rho correlation (p = -1.0; p = 0), this pattern is sta- nisms since both adult turtles, tortoises, and ele-
tistically highly significant. Therefore, the pres- phants have few natural predators. If an invasive
344 AMERICAN ANTIQUITY [Vol. 68, No. 2, 20031

Table 3. Clovis Assemblage Data Summarized by Body Size

Bison, equids, Other ungulates, sloths, Lagomorphs Rodents

Proboscideans and camelids and glyptodonts and armadillos and insectivores
Size Class 1 2 3 4 5
N Taxa 3 9 9 4 30
Total MNI 91 110 37 25 93
MNI per taxon 30.3 12.2 4.I 6.3 3.1

Expected n sitest 3.3 9.9 9.9 4.4 33

Actual n sites 26 19 18 9 14
Act. - Exp. n sites 22.7 9.1 8.1 4.6 -1 9
Note: tSee text for explanation of calculation.

species, in this case prehistoric humans in Pleistocene most recent foraging populations occupy primarily
North America, is expected to move into a niche with "marginal" environments and must coexist with
minimal interspecific competition, then the prey duo neighboring nonforaging societies, their subsistence
of elephants and tortoises is not unexpected. options are perhaps significantly more limited than
the constraints on hunter-gatherers of the past. Also,
Is Big-Game Hunting Possible? hunter-gatherers documented in the modern era
The Clovis faunal record certainly implies the exten- occupy ecosystems that have been inhabited by
sive and selective use of large-bodied prey. Although humans for thousands of years and they likely exist
potentially the result of biased archaeological visi- at relatively high population densities (as compared
bility and recovery, the distribution of Clovis faunal to Clovis). For Clovis populations living in late Pleis-
assemblages appears more congruent with a special- tocene North America, land-use options would have
ized hunting strategy. Some critics, however, claim had few limits since population densities would have
that specialized hunting was simply not a viable sub- been extremely low. In such a situation of relatively
sistence strategy for Clovis peoples. As previously few people and abundant big-game availability, hunt-
discussed, theoretical arguments launched against ing specialization may have been possible in envi-
Clovis-as-specialists frequently point to the fact that ronments where this strategy does not and could not
ethnographically known examples of foragers with exist today. In most contemporary environments,
specialized hunting economies are geographically large-game specialization is simply not possible
limited to arctic and grassland environments (Meltzer because there are too many people and/or too few
1988, 1993; Meltzer and Smith 1986). large animals, a situation that has existed for perhaps
The ethnographic record does evidence that a thousands of years.
strong relationship exists between the proportion of If big-game hunting specialization is a viable
subsistence resources derived from hunted game and strategy in any environment as long as human pop-
environmental context (Keeley 1988; Kelly 1995; ulation densities are low relative to available game,
Murdock 1981). With few exceptions, hunter-gath- then there is little reason to presume that Clovis peo-
erer groups who derive more than 50 percent of their ples could not have been specialists simply because
food resources from hunting generally occupy envi- they occupied an ecologically diverse environment.
ronments with low mean effective temperatures (i.e., To examine the relationship between reliance on
arctic ecosystems), in areas in which the bulk of pri- hunting, environment, and population density, a sam-
mary biomass cannot be consumed by humans but ple of 92 foraging groups was examined. Populations
is accessible to herbivores (i.e., grasslands and tun- of hunter-gatherers that derive all of their subsis-
dra), and less commonly, in tropical ecosystems tence resources from foraging were collected from
where foragers trade meat for plant products with the Atlas of World Cultures(Murdock 1981).The per-
local, nonforaging, peoples (reviewed in Kelly cent of the diet comprised of plantlsmall terrestrial
1995:66-73). But these patterns are drawn from fauna and larger game resources was tallied. In addi-
modern and historically known hunting and gather- tion, the environment occupied and population den-
ing groups, and do not necessarily reflect the range sity for each group was recorded (data from Kelly
of possible subsistence behaviors of the past. Since 1995: Table 6-4).

Size Class

Size Class

Figure 6. (a) (top) Prey size class versus total MNI per species, calculated as the sum of MNIs for all taxa in each size-class
grouping, divided by the total number of species in that group (two-tailedSpearman's p = -.9; p = .037); (b) (bottom) Prey-
size class vs. the actual minus the expected number of sites per taxonomic grouping, assuming a generalist subsistence strat-
egy (two-tailedSpearman's p = - 1 ; p = 0).

Plots of log population density (people per 100 ing than would be predicted by their population den-
km2) for the sampled hunter-gatherer groups and sities. Hunters of the Great Plains are an interesting
their reliance on both plants/small fauna and large exception because, as noted by Meltzer (1993), they
game are presented in Figure 7a and 7b. Not sur- were armed with rifles and had the use of the horse.
prisingly, dependence on plants and small fauna As predicted by the diet-breadth model, the intro-
increases with population density and is statistically duction of the horse and gun should have caused a
significant (Spearman's p = .397, p < .001). The contraction of diet breadth because greater mobility
average dependence on larger game decreases with would have increased encounter rates with large
greater population density for all environments mammals, notably bison, and rifles would have
(Spearman's p = -.68, p < .001), the only exceptions decreased handling costs. A similar effect was
being the Mbuti and groups of the North American observed with the introduction of the snowmobileto
Great Plains who both tend to depend more on hunt- the Cree (Winterhalder 1981).
AMERICAN ANTIQUITY pol. 68, No. 2,20031

' Dcscrt

Forc st


A Plains

Population Density (persons/100 km 2,

rr o o arr r x ex -
X m r r -m--- --

Population Density (persons/100 km 2,

Figure 7. (a) (top) Proportion of plants in the diet vs. population density for a global sample of hunter-gatherers (n = 92,
Spearman's p = 397,p c .001); (b) (bottom) proportion of large game in the diet vs. population density for the same groups
(Spearman's p = -.68,p < .001).

Dividing the sample into groups who derive more km2. These findings cannot directly implicate Clo-
than 46 percent of their subsistence by hunting from vis peoples as big-game hunters, but it does estab-
those who do not (Figure 8), a two-tailed t-test estab- lish that hunting specialization can and frequently
lishes significant differences between their mean does occur at low population densities.
population densities ( p < .001). The mean popula- A problem of equifinality arises when these results
tion density value for groups with greater than 46 are compared with the findings of others who have
percent reliance on hunting is .25 people per 100km2, correlated dietary hunting dependence with environ-
and for less than 46 percent it is 37.5 people per 100 mental parameters (Binford2001; Keeley 1988,1995;

We argue that large-game specialization is only

seen in recent times in areas where there are few other
subsistence options. The inability to switch subsis-
tence strategiesin the arctic, for example, 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, that
of top carnivore. In other biomes where large ani-
mals were readily available, humans chose to be
large-game specialists early on, but due to increas-
ing numbers of people and decreasing numbers of
prey, they moved into a more generalized subsis-
tence strategy, eventually allowing for higher human
population densities.Furthermore, although the envi-
ronments where large-game specializationoccurs in
recent times are characterized by relatively low pri-
mary production, a vast majority of the primary bio-
mass present is accessible to large ungulates (due to
the paucity of woody vegetation), allowing these ani-
mals to attain and maintain high population densi-
Percentage of Large Game in Diet ties. These factors, we suggest, have shaped the
modern ethnographic record of large-game hunting.
Figure 8. Percentage of large game in the diet vs. population Therefore, regardless of the diverse environments
density for a worldwide sample of hunter-gatherers (n = 92,
two-tailed t-test,p < .MI). Note Y-axisis the log,, of popula- they occupied, as long as Clovis peoples maintained
tion density. Data from Kelly (1995:Table 6-4) and Murdock low population densities and large game was avail-
(1981). able and regularly encountered,a hunting-based sub-
sistence economy seems highly plausible.
Kelly 1995; Meltzer 1993). In other words, three
independent variables are collinear (population den- Conclusions
sity, environment, and subsistence), resulting in a Although there has been much debateregarding Clo-
sticky problem of establishing causality. An argu- vis diet breadth and subsistence strategy, few
ment could be made that environment is the proxi- attempts have been made to quantitatively analyze
mate cause of subsistence practices and population Clovis faunal assemblages. The zooarchaeological
density. We argue above with respect to the diet- data collected for the current analysis are admittedly
breadth model that large-game specializationshould imperfect, but provides one important source of evi-
result from relatively high encounter rates with large dence for the examination of Clovis prey choice.
game. Therefore, subsistence should ultimately be a Many potential biases exist in the dataset. Differen-
product of the density of prey coupled with the den- tial recovery and preservation of animal remains that
sity of human hunters, independent of environmen- bias the record toward large animals are possible
tal parameters. Although prey densities will ultimately influenceson the Clovis record that currentlyremain
be controlled by environmental productivity, human difficult to assess. Nonetheless, for the purposes of
population densities can be modified by adjusting this study, while acknowledging the existence of
"territory" or range size. Absolute numbers of these nonrandom factors influencing the dataset, we
humans, however, should be limited by food sup- are not willing to assume that this bias is so exten-
plies, whichis ultimately limited by subsistenceprac- sive as to reverse our findings.
tices. That is, hunter-gatherers who depend entirely We have framed the contrast between hunting
on hunting will exist at far lower densities than those specialization and generalization on the degree of
who adopt a more "vegetarian" lifestyle since hunters selectivityexpressed in prey choice in amanner con-
only exploit a small fraction of the available edible sistent with optimal foraging theory. Selectivity has
biomass in an ecosystem. been defined as the relative degree to which certain
348 AMERICAN ANTIQUITY [Vol. 68, No. 2, 20031

taxa are not exploited upon encounter. Specialists erers show a clear increase in hunting of large ani-
tend to bypass numerous potential prey in favor of mals as human population densities decline. Since
higher-ranked resources. The quintessential gener- Clovis represents the colonization of an uninhabited
alist, on the other hand, would exploit all or nearly landscape or slightly postdates that event, early Pale-
all prey encountered.Despite their presentation here oindian population densities were undoubtedly low.
as alternative strategies, they are intended only to Low population densities coupled with frequent res-
describe two possible extremes of a continuum. idential mobility over large distances provide a con-
Based on the analysis of 33 faunal assemblages it text where large-game specialization is not only
has been argued that Clovis hunting behaviors appear possible but also likely. Following Kelly and Todd
more closely aligned with a specialized,rather than (1988), subsistence emphasizing large-game hunt-
generalized, strategy. ing is not restricted to any particular environment
This conclusion is drawn from the relative abun- (except those lacking large game), and is perhaps the
dances of a range of taxa represented in Clovis sites. ultimate strategy for maneuvering among diverse
Unlike the expectations for a generalist strategy, the environments.
taxa most frequently occurring in sites and occur-
ring in the greatest numbers are those that would have Acknowledgments. We would like to thank Mary Stiner, C.
been infrequently encountered by Clovis hunter- Vance Haynes, Robert Kelly, Mary Lou Larson, Marcel
gatherers. This does not mean that Clovis existed by Kornfeld, Paul Martin, David Overstreet, David Andersen,
Marvin Kay, and one anonymous reviewer for their helpful
mammoth alone. Nor does it imply that Clovis peo- comments and insightful suggestions regarding this work.
ples only hunted mammoth, bison, and other large Although some of these commenters ultimately disagree with
animals. There is clear evidence that they did exploit our interpretations, we value their patience and contribution to
small mammals and reptiles on occasion. However, our own thoughts regarding Clovis subsistence. Gratitude is
based on estimated encounter rates, Clovis hunter- also extended to Julio Betancourt and Maria Nieves Zedeiio
who translated the abstract, and to Jeffrey Saunders who gen-
gatherers often ignored opportunities to harvest erously allowed the use of his unpublished faunal inventories
smaller game species, likely in favor of obtaining a of the San Pedro Clovis sites. And a special thanks to Cherie
higher-ranked resource. Turtles and tortoises are one Freeman who helped compile data for this study, and has
clear example of small game taxa that Clovis for- always encouraged us to pursue our interests. For her enthusi-
agers may have regularly taken upon encounter since asm in volunteering and for her companionship, we are very
much indebted.
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2000 The Tortoise and the Hare: Small Game Use, the Broad Accepted September 6, 2002.

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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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