Settlement Patterns, SourceâSink Dynamics, and Artiodactyl Hunting in the Prehistoric U.S....

Settlement Patterns, Source–Sink Dynamics, and ArtiodactylHunting in the Prehistoric U.S. Southwest

Karen Gust Schollmeyer & Jonathan C. Driver

Published online: 25 October 2012# Springer Science+Business Media New York 2012

Abstract Numerous studies in the US Southwest suggest that prehistoric artiodactylpopulations in areas of dense human settlement experienced population reductionswhich substantially reduced their availability to human hunters. Although mostassemblages from villages in this region are dominated by lagomorphs, some settle-ments maintained greater access to artiodactyls. Factors influencing this variabilityinclude both local settlement history and settlement location relative to productivesource areas for large game. In our study areas, source–sink dynamics likely contrib-uted to the long-term resilience of hunted artiodactyl populations and allowedvillagers continued access to animals moving in from source areas despite relativelyrapid game depletion in heavily hunted areas immediately around villages.

Keywords Hunting . Zooarchaeology . Settlement patterns . US Southwest

The recursive relationship between human resource acquisition activities and the environ-ment in which those activities take place is an important issue both in archaeology and formodern conservation efforts. One aspect of this relationship, the effects of human huntingand anthropogenic landscape change on large mammalian game species, has been thesubject of much debate in recent years. Although there are a number of examples ofresource depression affecting human access to large mammal populations in both thearchaeology and conservation biology literature (e.g., Badenhorst and Driver 2009;Broughton et al. 2010; Fa et al. 2005; Grayson 1991; Jerozolimski and Peres 2003;Wolverton et al. 2008), there are also counterexamples (e.g., Butler and Campbell2004; Vickers 1991). The relationship between human activities and changes inaccess to large mammals at different spatiotemporal scales is highly complex. Here,we examine factors influencing this relationship in the US Southwest during theperiod from 840 to 1400 AD. We suggest that insights from biological studies of

J Archaeol Method Theory (2013) 20:448–478DOI 10.1007/s10816-012-9160-5

K. G. Schollmeyer : J. C. DriverDepartment of Archaeology, Simon Fraser University, Vancouver, BC, Canada

K. G. Schollmeyer (*)School of Human Evolution and Social Change, Arizona State University, Box 2402, Tempe, AZ85287-2402, USAe-mail: karen.schollmeyer@asu.edu

source–sink dynamics (e.g., Novaro 2004; Pulliam 1988) explain some of the patternswe see in prehistoric large game use.

Large mammals tend to be highly desirable resources for humans. Ethnographicstudies in the US Southwest have shown that large mammals are among the mostsocially valued foods, particularly relative to locally available small mammals(Beaglehole 1936; Parsons 1925; Potter 2004). In anthropology, models of dietbreadth and prey choice posit that resources are often ranked based on energy returnsand that large-bodied prey normally rank higher than small mammals in such asystem (e.g., Broughton et al. 2011; Grayson and Cannon 1999; Grimstead 2010;Lupo 2007; Ugan 2005; Winterhalder and Smith 2000). In such a model, foragersmaximizing return rates may pass over lower-ranked resources (such as small mam-mals) if their chances of encountering higher-ranked (often larger) prey are accept-able. Recent studies by conservation biologists of tropical hunting show a similarpattern among contemporary hunters (Jerozolimski and Peres 2003; Peres andNascimento 2006; Vickers 1991). Given this pattern of general preference for largemammals over small ones, a decrease in the abundance of large mammals relative tosmall ones in archaeological assemblages may be interpreted as reflecting a decreasein large mammal availability in the hunting area in many situations (Badenhorst andDriver 2009; Cannon 2000; Grayson 1991; Janetski 1997), although this is notalways the case (Broughton 2002; Dean 2007; Szuter and Bayham 1989).

The US Southwest was occupied in later prehistoric times by people whose diet wasdominated by domestic crops, notably maize, squash, and beans. They had no largedomestic animals, and the one domestic bird (turkey) was only abundant in compara-tively mesic habitats. Wild mammals were hunted and trapped, and there was asignificant dichotomy between relatively large-bodied artiodactyls, such as deer, prong-horn antelope, and bighorn sheep, and much smaller lagomorphs (jackrabbits andcottontails) and rodents, such as prairie dogs and ground squirrels. The typical settle-ment ranged from a few dozen to a few hundred people; settlements were residentiallyabandoned regularly, and regional human populations experienced cycles of increaseand decline. Although some Southwestern communities used pit houses, the communi-ties that we use as case studies built multi-roomed masonry structures. Even withoutexcavation, the number of rooms can often be estimated from surface characteristics,and room counts serve as a proxy for human population estimates.

Archaeologists in the US Southwest have long used the artiodactyl index, a measure oflarge mammal relative abundance, as a means of comparing assemblages in order to assessspatial differences in hunters’ use of large game or changes in that use over time andinferred differences in artiodactyl availability (Badenhorst and Driver 2009; Broughton1994; Cannon 2000; Grayson 1991; Szuter and Bayham 1989). Interestingly, theartiodactyl index is highly variable in assemblages associated with sedentary agricul-tural groups in the Southwest (Badenhorst and Driver 2009; Schollmeyer and Driver2012). Although lagomorphs and other small mammals are nearly always morecommon than artiodactyls in assemblages from Southwestern villages, the degree towhich they outnumber artiodactyls varies enormously across both time and space.Throughout the Southwest, regional and interregional studies show a broad pattern ofgenerally low artiodactyl relative abundance punctuated by assemblages with rela-tively more artiodactyls (Badenhorst and Driver 2009; Broughton et al. 2010; Dean2007; Schollmeyer and Driver 2012). What factors influence this pattern? Are certain

Settlement Patterns, Source–Sink Dynamics, Hunting in the U.S. Southwest 449

village characteristics regularly associated with relatively greater access to largemammals, as expressed in artiodactyl relative abundance? Here, we examine thisissue using three case studies: the Mimbres region from 1000 to 1200 AD, the Zuniregion from 1125 to 1400 AD, and the Mesa Verde region from 840 to 1285 AD.

Many factors could potentially influence artiodactyl relative abundance in assemb-lages from prehistoric agricultural villages whose inhabitants did not raise large domes-tic animals. Here, we focus on local human population size, village location in relation tosurrounding settlements, elevation, and local settlement and population history. In thesecase study regions, our results show little correspondence between local human popu-lation size and artiodactyl relative abundance. However, there are interesting differencesamong assemblages with differing levels of access to hinterland areas with a lowerhuman population, particularly when those hinterlands are relatively mesic upland areasrather than more arid lowlands. Settlement history is also an important variable; settle-ments in previously unoccupied or lightly populated areas had initially greater access toartiodactyls, but this effect does not seem to have lasted long. This study indicates thatartiodactyl relative abundance declines quite rapidly after the initial appearance ofsedentary agricultural villages in an area, with archaeologically visible impacts afterjust a few decades. These findings support conservation biology studies that documentthe rapidity of human impacts on mammalian prey species. Our results are alsoconsistent with a pattern in which lightly populated hinterlands serve as source areasfrom which game may move into heavily hunted areas around villages. Although workremains to be done on this idea, this study suggests that source–sink dynamics may haveplayed a key role in the long-term resilience of regional animal populations and thesustainability of hunting in the US Southwest.

Artiodactyls in Archaeological Assemblages: Potential Factors Contributingto Inter-assemblage Variability

A large number of factors have the potential to contribute to inter-assemblagevariability in the relative abundance of artiodactyls. These include both taphonomicissues related to the deposition of assemblages and non-taphonomic factors. There aretwo main taphonomic issues involved here. First, it is difficult to relate the relativeabundance of taxa in an archaeological assemblage to their absolute abundance inprehistoric diets. Taphonomic processes act differently on different taxa, and themixed, disarticulated, and fragmentary specimens we find archaeologically cannotbe translated reliably into a direct count of complete animals (Lyman 2008). Com-pounding this issue is a closed-array problem in which an increase in the relativeabundance of one taxon may reflect either an actual increase in its abundance or adecrease in the abundance of another taxon in the assemblage. A second potentialproblem is variation in the excavation, screening, and analysis and reporting techni-ques used by different zooarchaeologists (Driver 1992). In this analysis, we examinegeneral trends in large mammal relative abundance across many assemblages,an approach we assume will cancel out particularistic biases in taphonomy,sampling, and analysis associated with individual assemblages and will notsignificantly skew the broader trends we identify. We also include onlyscreened assemblages. Within each of the regions examined here, many of the

450 Schollmeyer and Driver

assemblages were identified by a small number of analysts using very similarmethods.

A number of factors may contribute to the variability in the relative abundance oflarge game in archaeological assemblages. If taphonomic factors are held constantand variability reflects actual differences in game acquisition, these potential influ-ences can be grouped into three general categories: non-anthropogenic environmentalfactors, anthropogenic influences on environments and game populations, and otherhuman social practices (Table 1). Here, we examine both spatial and temporalvariations in an attempt to tease apart some of these factors. Data are not availablefrom our case studies to control for all of the potential confounding factors listed here.However, using data from three case study areas and from multiple assemblageswithin each area helps us control for some of these factors and to examine others inmore detail.

Non-anthropogenic environmental factors influencing variability in artiodactylrelative abundance show important spatial and temporal variability in the Southwestand can be grouped into five general categories (Table 1). Habitat productivity forboth large and small game depends on factors such as the availability of water, foodavailability and nutritional content, cover, and pairing success for breeding animals(Pulliam 1996), all of which may vary across time and space. Temporal variation inhabitat productivity may be influenced by climate change (e.g., Bousman et al. 2002;Broughton et al. 2008; Byers et al. 2005; Grayson and Delpech 2005; Wolverton2005). In our case study areas, paleoclimatic reconstructions show no indications ofclimate changes that would have substantially changed landscape productivity at abroad scale for artiodactyls or lagomorphs during the period we consider (Cannon2001, pp. 207–256; Varien et al. 2007). Spatial differences in habitat productivity areaddressed in this study by examining elevation differences among sites within thecircumscribed case study regions. Although more subtle habitat productivity differ-ences in time and space are likely, they are beyond the scope of this paper.

Table 1 Major non-taphonomic factors influencing large game acquisition in the US Southwest

Non-anthropogenicenvironmental factors

Anthropogenic factors affectingenvironments and animal populations

Other humansocial practices

Habitat productivity for largeand small game

Regional human population density Hunting technologies

Reproductive potential of largeand small game species

Duration of contemporaneous andprevious human occupations

Hunting techniques(e.g., communal versusindividual)

Patches with dense concentrationsof small game

Density of contemporaneous andprevious human occupations

Restrictions on groupmembers allowed tohunt certain taxa

Spatial distribution of potentiallyproductive agricultural areas

Degree of settlement clusteringor dispersal

Management of wild game(e.g., deliberate feeding,hunting quotas)

Non-anthropogenic spatialdistribution of productive largeand small game habitat areas

Anthropogenic spatial distribution ofproductive large and small gamehabitat areas, including distancefrom settlement

Scheduling of otheractivities

Interspecies variation in the reproductive potential of the various Southwesterngame taxa are not an important variable in this analysis as the same taxa are comparedacross all assemblages. Dense concentrations of small prey may increase theirranking relative to larger game in a prey choice model if the cost of processing thesmall prey item is low (Ugan 2005 and references therein). Densely concentratedsmall game may also have been more heavily exploited if large animals were scarce,or if children or others socially prohibited from hunting large animals were involvedin collecting small game. Prairie dogs are the only Southwestern mammal taxonnaturally available in dense concentrations large enough to substantially affect theirrelative abundance in this way; we argue later that some communities may have hadaccess to large prairie dog towns based on their faunal assemblages, but the prehis-toric locations of these resources are difficult to pinpoint with the data currentlyavailable. Finally, in the absence of climatic changes, the distributions of potentiallyproductive agricultural areas and of animal habitat are unlikely to have changedover time due to non-anthropogenic factors, but they do show important spatialvariability in relation to the locations of the communities providing faunal assemb-lages. We approach this variability using faunal assemblages from agricultural vil-lages in a variety of locations. In each case study area, some villages examined herewere located in more densely settled areas, whereas others were nearer the edges of asettlement system and had greater access to more lightly populated hinterland areas.

Anthropogenic factors also vary across time and space. The anthropogenic factorsmost commonly identified as influences on artiodactyl relative abundance in theSouthwest are anthropogenic habitat change and artiodactyl population depressionor recovery, both of which are linked to the size of human populations and theirdegree of sedentism (Badenhorst and Driver 2009; Cannon 2000; Grayson 1991;Janetski 1997; Schollmeyer 2011; Szuter and Bayham 1989). Anthropogenic habitatchanges (particularly clearing land for fields and fuel collection and the planting offields and gardens) influence local landscape productivity for both large and smallmammals, reducing the availability of mature vegetation required by some specieswhile also adding attractive food sources and increasing the local availability ofdisturbance vegetation used by other taxa (Cannon 2000; Driver 2002; Driver andWoiderski 2008; Neusius 2008; Schollmeyer 2005). Resource depression may haveoccurred as increasing human populations placed greater demands on higher-rankedtaxa, which in the Southwest were likely large game species.

Attributes of human populations that influence anthropogenic habitat change andresource depression can be grouped into five general categories (Table 1). Regionalhuman population density influences both the demand for game animals and thedegree and extent of habitat change, as does the density and duration of localpopulations and settlements during both the time period from which an assemblageis drawn and previous occupations of the area. Settlement clustering or dispersal hasa high potential to impact both landscape productivity and the effects of humanhunting on animal populations. A number of contemporary studies indicate thatwhere villagers hunt, areas within 10 km or less of a village have substantially lowerdensities of large-bodied game taxa, including artiodactyls (Hart 2000; Hill andPadwe 2000; Mittermeier 1991; Peres and Nascimento 2006; Van Vliet et al.2010). A closely related attribute, the spatial distribution of productive habitat areasrelative to villages, would also have influenced access to game (Broughton 2002;

Dean 2007). Villages located near the edge of a settlement cluster near a more mesic,productive area for large mammals may have greater access to these animals thanthose located at the edge of a cluster in more xeric areas. In order to control for orexamine these factors, we selected case study areas and time periods with reliableregional- and site-level population estimates, well-understood settlement patterns,and precise dating of archaeological assemblages.

Other human social practices also have the potential to influence large game relativeabundance in the Southwest. Some of the issues raised by such practices are relatively easyto address, but others are quite difficult to examine archaeologically (Table 1). Changes inhunting technologies could increase the profitability of hunting small mammals, thuspotentially changing their ranking in prey choice models (Grayson and Cannon 1999;Hockett and Bicho 2000; Lupo and Schmitt 2002; Madsen and Schmitt 1998; Schmittet al. 2002). There is no evidence of the development of new hunting technologiesduring the time period examined here, as discussed below. However, it is possible thathunting techniques changed in popularity, particularly shifts in the frequencies ofindividual hunts relative to communal hunts such as jackrabbit net hunting that mighthave changed the relative abundances of these animals in assemblages. Such changescannot be entirely ruled out in the assemblages examined here; evidence for themvaries by case study area and is discussed in more detail below.

Shifts in social rules regarding group members allowed to hunt certain taxa couldalso influence the relative abundance of different types of game. Ethnographic studiesof traditional societies suggest that such rules are common; for example, rules mayrestrict the hunting of large game to adult men or other age- and gender-dependentcategories (Lupo and Schmitt 2002; Owen 2005; Szuter 2000). Unfortunately,changes in these rules may be difficult to distinguish in the archaeological record;none have been suggested for the study area. Management of wild game (such asdeliberate feeding, socially bounded hunting territories, or hunting quotas) is alsodifficult to distinguish archaeologically and has not been documented in our studyarea and time period. Finally, the scheduling demands of other activities may havelimited large game hunting in some times and places. In our study area, agriculturaltasks would have taken substantial amounts of time and required at least part of thepopulation to remain near fields at some times of year; in the Mesa Verde and Zuniregions, the demands of turkey husbandry would also have been a factor during sometime periods. In this study, agricultural products contributed the bulk of human dietsin all three case study areas throughout the time periods we examine, making differ-ences in the scheduling of agricultural tasks unlikely to have influenced inter-assemblage variability in the artiodactyl index. The importance of turkey husbandryvaries temporally and is discussed in more detail for the relevant assemblages.

In summary, this study attempts to hold a number of variables constant. We focuson comparing assemblages with differences in local human population size, locationin a settlement system (e.g., at the interior or the edge of a settlement cluster),elevation, and local settlement and population history. The variables we have chosenare particularly important in two types of explanations for changes in large mammalrelative abundance: resource depression and source–sink dynamics. Resource depres-sion, as discussed above, is the idea that increasing human populations and/orincreasing sedentism lead to increased consumption of higher-ranked resources likelarge mammals. Overharvesting leads to declines in their abundance, which in turn

influences hunters to add previously neglected lower-ranked resources to the diet.The variables of human population size and local settlement and population historyare both important in this model.

Source–Sink Dynamics and Access to Game Animals

Source–sink dynamics are not discussed widely in the archaeological literature, butprovide a promising avenue of research to explain some types of variability in large gameaccess. In conservation biology, a number of studies indicate that less heavily hunted areas,particularly those with high productive potential for game species, may serve as sourceareas for game animals captured in heavily hunted sinks around human settlements.Sources are populations or subpopulations of a given taxon in which births exceed deathsand emigration exceeds immigration (Novaro 2004; Pulliam 1988). Source areas maysupply surplus individuals to sinks, defined as populations or subpopulations inwhich deaths exceed births, immigration exceeds emigration, and game populationswould decline and vanish without source populations to replenish them. Whether aparticular area is a source or a sink depends on the characteristics of its habitat. Somehabitats are more suitable for the survival and reproduction of a taxon than others,either because of differences in the availability and quality of resources (food, water,nesting sites) required by that taxon or because of differences in mortality rates (fromhuman hunting or other predation, disease, or other factors; Pulliam 1996). Whenmigration between two habitats is possible, the more productive one may become asource area from which individuals move to the less productive sink area.

In studies of hunting, the distance of an area from human settlements or other easyaccess points (such as roads) is often one of the primary factors influencing whetheran area is a source or a sink for a given taxon. Many of these studies are carried out incomparatively homogeneous tropical and neotropical environments in which otherhabitat differences are minor. These studies often indicate that areas within 10 km orless of a village are game sinks for many taxa, while surrounding unhunted or lightlyhunted areas act as sources. Studies of Ache hunters in Paraguay show that althoughthe densities of most prey species are substantially lower in hunted areas than inprotected reserves, areas more than 6 km from a village or point of hunter entry intothe reserve have prey animal densities similar to unhunted areas deep within it (Hilland Padwe 2000; Hill et al. 1997). Other studies in tropical and neotropical forestsshow similar patterns (Hart 2000; Mittermeier 1991; Peres and Nascimento 2006).Although distance from settlements often dictates the boundaries between sourcesand sinks in these studies, sources may also be defined by culturally created bound-aries. Such areas are particularly important in preserving source populations ofhunted animals where the hunting areas of multiple settlements overlap or wherevehicular travel is common. In a large-scale study of African bushmeat hunting, Fa etal. (2006) found a significant inverse relationship between distance to a protectednational park and captured meat per person across both villages and urban centers,suggesting that the parks are game sources for their entire surrounding regions.Prehistorically, it is possible that sparsely populated territories separating mutuallyhostile cultural groups created socially bounded spaces that also protected gamepopulations (Driver 2010; Kay 1994; Martin and Szuter 1999).

Anthropogenic activities may create “traps” or patches of habitat that appear suitable fora species’ survival and reproduction, but which actually are unsuitable because of character-istics immigrating animals cannot detect (Pulliam 1996). Mechanically tilled agriculturalfields, for example, often have plant species and other characteristics used by nestingbirds as indicators of suitable habitat, but which in fact are unsuitable nesting sites asa result of annual disturbance from farm machinery late in the nesting cycle (Pulliam1996). Gardens, fields of all kinds, and orchards may also be “traps” for animalsattracted to disturbance vegetation because frequent opportunistic hunting in theseareas by agriculturalists in the course of other activities makes game taxon mortalityrelatively high (Smith 2005). Although fast-reproducing “garden game” species maydo well under such conditions (Daily et al. 2003; Linares 1976; Naughton-Treves2002; Peres 2001; Smith 2005; Stahl 2008), slower-reproducing species that favorsimilar environments may in effect mistake these areas for source habitats eventhough hunting or other types of disturbance make them sinks for that species.

We suggest that source–sink dynamics may have contributed to some of the patternswe identify in artiodactyl relative abundance in archaeological assemblages, particularlythe differences among contemporaneous settlements located in different environmentsand locations within a settlement system. We expect that most hunting in all of thevillages we examine took place near the village and that villages located close to sourceareas had more access to artiodactyls moving into their heavily hunted surroundingsthan villages located in other areas. This possibility is discussed in more detail below.

Methods for Comparing Archaeological Faunal Assemblages

In order to compare artiodactyl relative abundance among different assemblages, we use theartiodactyl index, the ratio of artiodactyls (measured as number of identified specimens, orNISP, identified to order or lower taxonomic levels) relative to the total NISP of artiodactylsplus lagomorphs (Broughton 1994; Cannon 2000; Driver 2002; Spielmann andAngstadt-Leto 1996; Szuter and Bayham 1989). Using this index avoids conflatingchanges in the use of mammal taxa less likely to represent human dietary resources(such as carnivores and some rodents) with changes in the relative abundance of foodtaxa, and it also avoids problems associated with comparing taxa identified todifferent levels, such as species with size class (Grayson 1991). We use NISP ratherthan MNI (minimum number of individuals) because the former is commonlyreported and does not vary between analysts in the way it is calculated (Grayson1984, pp. 28–40), an important consideration when comparing assemblages recordedby a number of different analysts who may have used somewhat different procedures.

The artiodactyl index was defined by Szuter and Bayham (1989, p. 84) as “theartiodactyl NISP divided by the sum of the artiodactyl and the lagomorph NISPs.”Artiodactyls are the only large mammals hunted for food in the Southwest. Lago-morphs are ubiquitous small game genera that are unlikely to be found as non-culturalintrusions in archaeological sites. Therefore, the artiodactyl index is a measure of therelative representation of large game in an assemblage. Specimens were included inthe artiodactyl count if they were identified to the level of the order (e.g. medium-sized artiodactyl), family, genus, or species. Similarly, lagomorphs include specimensidentified to order or family, as well as to genus or species.

In the Southwest, native Holocene artiodactyl species include deer (Odocoileus sp.) andpronghorn antelope (Antilocapra americana), both of which are commonly identifiedin archaeological assemblages. Elk (Cervus elaphus) and bighorn sheep (Ovis cana-densis) are sometimes identified in archaeological assemblages, but are rare in theregions and time period considered here. Native lagomorphs include jackrabbits(Lepus sp.) and cottontail (Sylvilagus sp.); pika (Ochotona sp.) are native to higherelevations, but have not been identified in any of the assemblages considered here.We also consider prairie dogs (Cynomys sp.) and turkeys (Meleagris gallopavo).

Local human population size is compared in several different ways. We use roomcounts from survey and excavation data as an index of local human population. In thethree case study areas, vegetation and geologic conditions are such that surfaceestimates of room counts are fairly reliable; although these counts are probably notabsolutely correct, they are adequate for determining the relative size of site or areapopulations in relation to each other. Room counts are used to estimate both therelative population of individual villages and the relative number of people using anarea. As discussed above, contemporary studies indicate that the majority of huntingaround sedentary villages occurs <10 km from the village; a number of Southwesternarchaeological studies use values of 6 km (Varien 1999). Here, we use a fairlygenerous estimate of 7 km as the radius within which most daily hunting takes place.The number of rooms within 7 km of a village (including rooms in that village andany neighboring settlements within that radius) serves as a measure of how manypeople were using a hunting area.

Case Study Area Analyses and Results

All three case study areas (Fig. 1) have well-established chronologies, well-understood settlement patterns and settlement histories, and high-quality faunal data,the latter of which is publicly accessible online in the Digital Archaeological Record(tDAR) database (http://www.core.tdar.org); we also used additional Mimbres data

Fig. 1 The three case study regions in the US Southwest

from published sources. Supporting data for individual sites are found in reports intDAR, on the Crow Canyon Archaeological Center web site (http://crowcanyon.org/research/research_publications.asp), and in various published reports. All three regionshave excellent survey coverage in the subareas discussed here such that the locations ofall sites with over 100 rooms and all or nearly all sites with over 40 rooms are known.Settlement data for the Mimbres and Zuni regions are drawn from a database compiledby Matthew Peeples. Settlement data for the Mesa Verde region is drawn from theVillage Ecodynamics Project database (Ortman et al. 2007; Varien et al. 2007).

Each of the three case study areas allows us to hold certain variables relativelyconstant and focus on others. The Mimbres and Zuni cases have somewhat broaderspatial and temporal resolution and most clearly show the effects of human population,elevation, and location at the interior or edge of a settlement cluster; the Zuni case alsohighlights the effects of population history. The Mesa Verde case is somewhat different,with greater spatial clustering of assemblages and more precise temporal resolution.Unlike the other two cases, the Mesa Verde data highlight the rapidity of changes inartiodactyl relative abundance and, by inference, local artiodactyl availability.

Elevation and Population: The Mimbres Region, AD1000–1200

The Mimbres region provides an example of spatial and temporal variability at afairly broad scale. Faunal assemblages examined here are from two locales, theMimbres Valley and the eastern Mimbres area, separated by the Black RangeMountains. Residents of this area lived in agricultural settlements by the latter partof the Late Pithouse Period (AD550–1000), and villages of up to several hundredrooms stood along floodplains during the Classic Mimbres Period (AD1000–1130).After 1130 AD, many of the villages were abandoned; some areas were depopulated,whereas others saw continuing occupation in smaller, scattered agricultural hamletsalong the same drainages during the Reorganization phase (AD1130–1200). Datingin the Mimbres area is not normally precise enough to assign sites or faunalassemblages to subdivisions within these temporal phases. We examine 15 assemb-lages in this study: five from the Classic Mimbres Period in the Mimbres Valley, fourClassic Mimbres assemblages from the eastern Mimbres area, and six Reorganizationphase assemblages from the eastern Mimbres area (Appendix 1).

Taphonomic factors are controlled in several ways. Assemblages from the Mim-bres Valley were excavated and analyzed by different projects and analysts, but allwere screened through a 0.25-in. mesh. In the eastern Mimbres area, all assemblageswere excavated by the same project with the same methods and were screenedthrough 0.0625-in. mesh; they were examined by three different analysts usingsimilar techniques. In both areas, faunal assemblages came from habitation room fill.Although different rooms were subjected to different taphonomic processes, the rangeof processes is expected to have been very similar across the archaeological sites.Environmental variables likely to be of importance include elevation differences andinterregional differences in aridity. The Mimbres Valley receives about 120 mm moreprecipitation than the eastern Mimbres area annually (Western Regional ClimateCenter 2005), and the Mimbres River flows perennially at the elevations of the sites

examined here. The eastern Mimbres area lies in a rain shadow formed by theintervening mountains, and the drainages examined there run only intermittently. Inboth areas, higher-elevation sites would have been relatively more mesic than those atlower elevations, with accompanying differences in vegetation and environmentalproductivity for game.

Hunting technique popularity, particularly the frequency of jackrabbit drives, isunlikely to have changed substantially during the time periods examined here.Several lines of evidence indicate that rabbit drives took place, including occasionaldepictions on Mimbres Black-on-White bowls from the Classic Period (Shaffer andGardner 1997) and evidence from at least one faunal assemblage (Schmidt 1999).Snares are also occasionally depicted on Classic Mimbres Black-on-White bowls(Shaffer et al. 1996). However, there is no evidence that the frequency with whichthese techniques were used changed over time (Cannon 2001, pp. 207–256).

Until recently, archaeologists speculated that the meat demands of the largepopulation of the Classic Mimbres Period led to declines in artiodactyl abundancethat contributed significantly to the abandonment of the Classic villages around AD1130 (Minnis 1985; Powell 1977; Shaffer and Schick 1995). However, more recentexaminations indicate that declines in large mammal availability occurred substan-tially earlier (Cannon 2000; Schollmeyer 2011). Throughout this region, relativelylarge sedentary populations were consuming local artiodactyls and altering the localenvironment by harvesting fuel wood and planting extensive agricultural fields wellbefore the assemblages considered here were deposited. Many or most Classic Periodvillages were constructed directly on Late Pithouse villages from the precedingperiod, so these preexisting impacts would have affected the surrounding resource-collecting areas. Thus, it is unlikely that anthropogenic habitat change explains thedifferences between the Classic Period assemblages examined in this paper.

In the eastern Mimbres area, access to artiodactyls did not improve during theReorganization phase, when groups dispersed into smaller settlements (Schollmeyer2011; Schollmeyer and Coltrain 2010). Instead, demand for large mammals likely farexceeded their local availability in all periods. However, isotope analyses suggest thatat least some of the deer that were acquired were local; some animals appear to haveeaten moderate quantities of maize, presumably from fields (Schollmeyer andColtrain 2010). In the Mimbres area, the most likely explanations for differencesbetween assemblages in large mammal relative abundance are spatial: site locationrelative to high-elevation areas and relative to the land use areas of other settlements.

The Mimbres region had a substantial sedentary agricultural population through-out the study area both before and throughout the periods examined here. Thus, ouranalysis focuses on population (site size in rooms and population in rooms within7 km) and elevation-linked environmental differences rather than on temporal changeor differences related to population history. There is marked variability in artiodactylrelative abundance in the Mimbres assemblages (Fig. 2 and Appendix 1). Notsurprisingly, lagomorphs outnumber artiodactyls by a large margin in every assem-blage. However, some assemblages have much lower artiodactyl indices than others.

We examine the effects of population size first. Local population size during theClassic Mimbres Period and Reorganization phase shows no strong correlation with

the artiodactyl index (Figs. 3 and 4). There is no significant correlation (r200.04)between site size or site population (as estimated by number of rooms) and theartiodactyl index value. There is also no correlation (r200.00) between the numberof people hunting within the 7-km area most frequently used by site residents (asestimated by room count within a 7-km radius) and the artiodactyl index. Dividingthe sites by subarea within the region and/or by time period does not substantiallychange this pattern for most site subgroups. When the Classic Period Mimbres Valleysites are considered alone, there is a significant relationship between the artiodactylindex and site room count, but not room count within 7 km. However, we attribute

Fig. 2 Composition of Mimbres area site assemblages

R²= 0.04310.00

Site room count0 50 100 150 200

Fig. 3 Mimbres site assemblage artiodactyl index by site room count

this pattern to the very strong relationship between the artiodactyl index and elevation(discussed below) as the sites with the highest number of rooms in this subgroup arethose located at the lowest elevations. This lack of a consistent relationship betweensite and local population sizes and the artiodactyl index is consistent with the idea thathuman impacts on local game populations predated the Classic Mimbres Period, asdiscussed above. A combination of anthropogenic habitat changes and prolongedhuman hunting pressure probably reduced local artiodactyl populations in the mostheavily used areas around all of these villages well before the assemblages weexamine here were deposited. This may have happened somewhat more slowly insmaller settlements and those with a lower nearby population, but by the time theassemblages we examine were deposited, these processes had already occurred.

Elevation is more closely correlated with the artiodactyl index. Initially, thisrelationship appears statistically significant, but somewhat weak (r200.13).However, the outlier visible in Fig. 5 has a great effect on the relationship. Thissite, Phelps hamlet, is at the upper edge of the settlement system in theReorganization phase, although its elevation relative to the Classic Period sites isonly moderate. When this site is removed, the correlation improves substantially (r200.42). Dividing the sites by subarea within the region and/or by time period also

R²= 0.001

Rooms within 7km radius0 200 400 600 800 1000 1200

Fig. 4 Mimbres site assemblage artiodactyl index by room count within a 7-km radius

R²= 0.1251

Mean elevation within 7km radius1400 1500 1600 1700 1800 1900 2000 2100

Fig. 5 Mimbes site assemblage artiodactyl index by elevation. Note that Phelps hamlet (the outlier) is atthe upstream end of the Middle Palomas settlement cluster in Fig. 2

produces consistently significant relationships between elevation and artiodactylindex. This underscores the idea that both elevation and settlement location areimportant. Sites at low elevations have relatively low artiodactyl index values, evenif they are at the lower edge of the settlement group (Fig. 2). Sites at higherelevations tend to have relatively higher artiodactyl index values, particularlyif they are also at the edge of a settlement group. This applies to several of thesites with the highest index values in their time periods, including both Phelpsin the Reorganization phase and Jackson Fraction in the Classic MimbresPeriod, located at a high elevation on an isolated side drainage in theMimbres Valley. Previous researchers have also noted positive relationshipsbetween large mammal relative abundance and elevation in the Mimbresregion (Broughton et al. 2010) and elsewhere (Szuter and Bayham 1989).

The effect of elevation appears to be independent of site population or the localpopulation around sites. We suggest that this pattern of lower large mammal relativeabundance in assemblages from lower-elevation sites reflects the fact that these morexeric areas were not highly productive for artiodactyls, even habitats outside theregular hunting areas of villages. Because of this lower productivity, lower-elevationareas did not act as sources from which substantial numbers of animals moved intothe more heavily hunted “sinks” near villages. In contrast, higher-elevation areaswere more productive for artiodactyls (particularly deer). In these areas, sites locatedadjacent to more sparsely populated source areas (such as Jackson Fraction in theClassic Mimbres Period and Phelps in the Reorganization phase) had relativelyhigher access to artiodactyls. The excellent survey data for this region allows us toassume that areas lacking residential archaeological sites had little or no residentialoccupation. In particular, the higher-elevation upland areas lacking residential siteswere largely unsuitable for maize agriculture. These areas would have been excellentpotential source areas for game populations. The relatively high artiodactyl indicesseen in assemblages from sites at higher elevations and at the edges of a settlementsystem are consistent with the idea that in the upland portions of the study area, deerhabitat outside the daily use areas of villages appears to have acted as a source areafrom which animals moved into the “sinks” around villages.

Elevation, Population, and Settlement History: The Zuni Region, AD1125–1400

Faunal data from the Zuni region provide a somewhat finer temporal and spatial scaleat which to examine variability in artiodactyl relative abundance. Although the regionsaw continuous occupation throughout the Prehistoric Period, there was some tem-poral variation in local settlement patterns as the area’s residents gradually movedupstream over time (Kintigh 1996). Around AD1200, this gradual movement under-went a more rapid pulse, with substantial movement into the El Morro Valley(Schachner 2007). Dating in this region is relatively precise, with assemblagesassigned to periods of 50–150 years. We examine 14 assemblages from four subareaswithin this region (Appendix 1). The Pescado and El Morro areas to the north havethe largest number of sites and the highest room counts. Populations were somewhatlower in the Ojo Bonito area and lowest in the Upper Little Colorado area to the

south. Assemblages come from the Pueblo III (AD1175–1275) and Pueblo IV (AD1275–1400) Periods, although some are dated more precisely within these periods.

The assemblages were excavated under a series of archaeological projectsdirected by the same researchers and using the same methods, includingscreening through a 0.25-in. mesh. The range of processes affecting the depos-its is expected to have been very similar across the archaeological sites exam-ined here, although the specific processes affecting the room fill and middenfill in any specific part of a site may have varied. Analysis of the fauna wasdone by three analysts using similar methods (Clark 1997; Potter 1997; Thiel2010).

Several previous zooarchaeological studies in the area have focused largely onother time periods or on research questions unrelated to large mammal relativeabundance (Clark 1997; Tarcan 2005). Potter (1997) focused on the El Morro Valleyand evidence for changes related to feasting practices accompanying the transition fromthe loose clusters of room blocks that comprised the area’s Pueblo III Period villages to thevery large, single-building, plaza-oriented pueblos of the Pueblo IV Period. His studyindicates that turkey production increased during this transition, a shift he argues waslargely the result of increased demands for dependable supplies of meat for communalfeasting. He suggests that communal hunting (for artiodactyls and lagomorphs) and long-distance hunting (for pronghorn) became more important over time, but he does notdocument any changes in artiodactyl availability. He suggests that the Zuni Mountainsjust northeast of the El Morro Valley remained a productive and reliable hunting ground.

Fig 6 Composition of Zuni area site assemblages

Like the Mimbres area, several of the most likely explanations for inter-assemblage variability in large mammal relative abundance are related to spatialvariables. Site location relative to high-elevation areas is likely to be of importance,as is site location relative to the land use areas of other settlements. Village populationand the population within 7 km of each site may also have influenced large mammalrelative abundance. Unlike the Mimbres case study, local population history is alsolikely to have influenced differences in large mammal relative abundance. Settle-ments in areas with earlier occupations may have had less access to deer by the timethe assemblages examined here were formed, whereas those in previously lightlyoccupied areas may have had access to relatively larger local artiodactyl populations.

In the Zuni area, our analysis focuses on population (site size and populationwithin 7 km) and elevation-linked environmental differences (Fig. 6) along withvariability in settlement history. As with the Mimbres case, local population sizeduring the Pueblo III and IV Periods does not seem to have greatly influencedartiodactyl relative abundance in the assemblages examined here. There is no corre-lation between site size or site population (as estimated by the number of rooms) andthe artiodactyl index (r200; Fig. 7). There is a slight but not statistically significantpositive relationship between the number of people hunting within the 7-km area

R²= 0.0001

Room count0 200 400 600 800 1000

Fig. 7 Zuni site assemblage artiodactyl index by site room count

R²= 0.024

Rooms within 7km radius0 500 1000 1500 2000 2500 3000 3500 4000

Fig. 8 Zuni site assemblage artiodactyl index by room count within a 7-km radius

most likely used by site residents (as estimated by room count within 7 km) and theartiodactyl index (r200.02; Fig. 8), the opposite of what one would expect if localpopulation demands were negatively influencing artiodactyl relative abundance bycausing local resource depression.

The most interesting picture in the Zuni area is gained by examining the artiodactylindex in site assemblages in light of three combined variables: elevation, positionwithin or at the edge of a settlement system, and settlement history. Although there isa significant correlation between elevation and the artiodactyl index (r200.24; Fig. 9),a better understanding of the relationship is offered by considering it in combinationwith settlement position within a cluster and settlement history. The sites with thelowest artiodactyl index values in their faunal assemblages (0.02 or below) are alllocated at elevations below 2,000 ft: Hinkson Ranch, Jaralosa Pueblo, and Ojo Bonito(Fig. 6 and Appendix 1). They are also located in the interior of a settlement clusterand in an area with a history of earlier settlement (Huntley and Kintigh 2004; Kintigh1996). This combination of factors severely limits the degree to which outside areascould act as a source for large game; these sink areas were located in a relatively lessproductive environment and were difficult for immigrating artiodactyls to reach. Theother two sites located below 2,000 ft—Baca Pueblo and Rattlesnake Point Pueblo—are located in an area with much wider spacing between settlements; their artiodactylindex values are relatively moderate.

Techado Spring, the site with the highest artiodactyl index value for itsfaunal assemblage, is at a relatively high elevation. It is also fairly isolated,at the edge of the settlement area. The El Morro Valley sites are also charac-terized by relatively high artiodactyl indices. As with Techado Spring, thesesites are located at high elevations, but they lie in an area of settlementsdensely clustered along one drainage area. Importantly, however, the El MorroValley was empty of permanent villages until this settlement cluster wasestablished around AD1250–1300 (Huntley and Kintigh 2004; Kintigh 1996).During the relatively short period in which these villages were occupied, the combi-nation of a lack of anthropogenic impacts from previous populations and a higher-elevation location in an area productive for deer appears to have acted to maintainrelatively high access to artiodactyls.

R²= 0.2438

1800 1900 2000 2100 2200 2300 2400 2500

Mean elevation within a 7km radius

Fig. 9 Zuni site assemblage artiodactyl index by elevation

Prairie dog use appears to have been quite patchy in this study area. The use of prairiedogs shows no particular pattern in terms of site size, elevation, or population history.Presumably, the presence or absence of substantial prairie dog towns near villagesdetermined how common they were in these assemblages, but it is difficult to determinewhat influenced the presence of prairie dog towns with the data available here.

Rapid Large Mammal Depletion with Relative Environmental Homogeneity:The Mesa Verde Region, AD840–1285

Faunal data from the Mesa Verde region provide unusually fine temporal and spatialresolutions. In this area, many assemblages can be assigned to 40-year time periods as aresult of numerous dendrochronological samples. This temporal resolution, combinedwith the availability of data from some of the earliest settlements in the local sequencethrough the end of Puebloan occupation in the area, allows us to examine some of thetrends identified in the other two case studies at a much greater level of detail.

According to Varien et al. (2007), an initial pulse of occupation by year-roundagriculturalists occurred around AD600–725, with an additional increase in occupa-tion (probably linked to immigration) around AD800– 840. This population shiftedfrom dispersed settlements into large villages around AD840–880. The period from880 to 1060 AD was characterized by emigration and low population, followed byanother episode of immigration between 1060 and 1100 AD. Population peakedduring the AD1225–1260 period, which also saw the construction of large commu-nity centers. A final emigration episode began around AD1260, and the region wasdepopulated by around AD1280. In this study, we examine 13 assemblages fromeight sites dated to five time periods: one from the AD840–880 (Late Pueblo I)coalescence into villages; two from the AD920–1060 (Early Pueblo II) period ofrelatively low regional population; three from the AD1060–1140 (Late Pueblo II)period of population growth; two from the AD1140–1225 (Early Pueblo III) period ofsustained high population; and five from the final period of dense occupation fromAD1225–1280 (Late Pueblo III; Appendix 1). The presence of well-dated depositsfrom multiple time periods at the same sites allows us to examine temporal changeswhile controlling for spatial variables.

The assemblages examined here were all excavated using similar methods, includ-ing 0.25-in. mesh screening. As in the other case studies, although the specificprocesses affecting the room fill and midden fill in any specific part of a site mayhave varied, the range of processes affecting the deposits is expected to have beensimilar across the sites. Driver and his students analyzed the fauna using consistentmethods (Badenhorst 2008; Driver 2000; Muir 1999; Rawlings 2006).

Previous studies have documented lower artiodactyl relative abundance in assemblagesfrom the Pueblo III period (AD1140–1300) in comparison to earlier assemblages. Thispattern has generally been attributed to a combination of reduced artiodactyl availabilitynear villages caused by local hunting and anthropogenic habitat change, an increasedemphasis on turkey husbandry, and constraints on the use of “wild” environments fartherfrom villages (Driver 2002). Although several studies of individual sites and broadregional analyses have been published, inter-assemblage comparisons of spatial

factors over a smaller area have received less attention. Site location relative to theland use areas of other settlements is likely to be of importance, as is villagepopulation and the population within 7 km of each site. Elevation differences arenot marked in the study area and are less likely than in the other case studies to be ofimportance as deer habitat and refuge areas with differential village access.

In the Mesa Verde area, our analysis focuses on population and settlementhistory. The villages considered here are all located within 25 km of oneanother, and there is little variability in elevation likely to have contributed todifferences in source area productivity. Instead, this case offers a fine spatialand temporal scale of resolution at which to examine the rapidity with whichartiodactyl resource depression can occur.

Our earliest assemblage is from the late Pueblo I Period (AD840–880), whenpopulations in the area first began aggregating into large villages (Varien et al. 2007).This assemblage has an artiodactyl index value of 0.15, most likely a function of thelack of preexisting human impacts on artiodactyl populations (Fig. 10). Two assemb-lages from the subsequent early Pueblo II period (assemblages from AD920–980 andfrom AD1020–1060) have similar or higher artiodactyl indices (0.13 and 0.25,respectively; Fig. 11). Varien et al. (2007) characterize this as a period of relativelylow population.

In the Late Pueblo II Period, assemblages look quite different. The three assemb-lages from the AD1060–1140 period have artiodactyl index values of 0.10 and below(Fig. 12), likely a result of a combination of the accumulating effects of humanhunting and anthropogenic landscape change over the preceding periods and anincreasing regional population. This shift must have taken place fairly rapidly in

Fig. 10 Composition of Mesa Verde area site assemblages, Late Pueblo I Period

order to make assemblages from this 80-year time period look so consistentlydifferent from those of the Early Pueblo II Period. Turkey representation also sees asudden increase in this period. Turkey domestication took place at this time in the

Fig. 11 Composition of Mesa Verde area site assemblages, Early Pueblo II Period

Fig. 12 Composition of Mesa Verde area site assemblages, Late Pueblo II Period

Mesa Verde area (Rawlings and Driver 2010); it is interesting that this appears to haveoccurred just as access to large mammals underwent a substantial decline. Some

Fig. 13 Composition of Mesa Verde area site assemblages, Early Pueblo III Period

Fig. 14 Composition of Mesa Verde area site assemblages, Late Pueblo III Period

researchers have suggested that turkey domestication was a response to decliningavailability of protein and fat from large mammals (Badenhorst and Driver 2009;Spielmann and Angstadt-Leto 1996), and the evidence from these assemblages lendssupport to that interpretation.

Assemblages from the Early Pueblo III Period (AD1140–1225) reveal anoth-er consistent decline in the artiodactyl index relative to all earlier periods (withindex values of 0.06 and 0.03; Fig. 13), inferred to show a sharp reduction in theavailability of large mammals. Human populations during this period were increasingtoward their peak levels, and the effects of this and earlier occupations within thesmall region continued to accumulate. The relative abundance of turkeys also in-creased substantially. By the Late Pueblo III Period (AD1225–1280), regional humanpopulations reached their highest levels in all of prehistory (Fig. 14). Four assemb-lages from this period have uniformly lower artiodactyl index values than those fromall earlier periods (0.04 and below) and high turkey relative abundance. As popula-tions reached their peaks, access to artiodactyls reached an all-time low, and the useof turkeys remained high in order to supply villagers with meat. Recent researchindicates that these turkeys were provisioned with maize (Rawlings and Driver 2010),which may have placed a substantial burden on local people to grow crops to feedboth themselves and their food animals.

One additional Late Pueblo III Period assemblage, from Yellow Jacket Pueblo, hasa substantially higher artiodactyl index value (0.21). There are two possible explan-ations for this. One is that this assemblage contains a mix of fauna from two differentuses of the site. At Sand Canyon Pueblo, Kuckelman (2010) demonstrated thatalthough secondary refuse (the vast majority of the Late Pueblo III assemblage)contained very few artiodactyl remains, the more limited immediate pre-abandonment contexts on the site had an artiodactyl index twice as high. Conversely,turkey remains were extremely common in secondary refuse and relatively rare inimmediate pre-abandonment contexts. She attributes this to maize crop failure shortlybefore the site’s abandonment, which forced people to increase their use of wildresources, probably at the cost of increased competition and conflict with neighboringpeople (Kuckelman 2010). When secondary refuse and abandonment contexts arecombined, artiodactyl indices are artificially inflated, and it appears as thoughartiodactyl access was higher throughout the Late Pueblo III Period. Ongoing exam-ination of the Yellow Jacket Pueblo assemblage will confirm whether a similar patternexists there.

A second cause of relatively higher artiodactyl values may relate to ancientpatterns of deposition and the sampling decisions of archaeologists. At bothSand Canyon and Yellow Jacket, artiodactyl remains are concentrated in depos-its associated with masonry tower complexes (Muir and Driver 2002); at SandCanyon, those same areas contain many abandonment-context deposits, and this mayalso be the case at Yellow Jacket. Alternatively, if the relatively high artiodactyl indexis indeed characteristic of the entire assemblage rather than limited abandonmentcontexts, it is also possible that Yellow Jacket’s position near the edge of thissettlement area, with fewer neighboring sites and greater access to more sparselypopulated surrounding land, allowed greater access to large mammals. This may have

been linked to the type of source–sink dynamics discussed for the Mimbres and Zuniareas.

As in the Zuni case, prairie dog use appears to have been spatially patchy in theMesa Verde region. It is difficult to determine what influenced the presence of prairiedog towns with the data available here, but these animals appear to have been quiteimportant in those areas in which people had reliable access to them.

Patterns in Case Studies and Implications for Future Research

Together, these three case studies highlight a number of interesting patterns. In areasin which large, sedentary human populations had been established for a century ormore before the assemblages we examine here were deposited, large mammal relativeabundance is quite variable. Both elevation and position at the edge versus the interiorof a settlement cluster appear to influence large mammal relative abundance in thissituation. We suggest that this pattern is related to source–sink dynamics. Lower-elevation areas were generally more xeric and less productive for large mammals, anddid not act as productive sources from which the game sinks around villages in theseenvironments could be refilled. Higher-elevation areas had greater productive poten-tial. We suggest that in areas with a long history of dense sedentary human popula-tions, local large mammal populations within about a day’s hunting distance of all thevillages had been depleted; every village’s regular hunting area was probably a “sink”for large game. Differences in large game access among these villages are attributablein part to their position relative to productive source locales from which animalsmoved into hunting areas. Villages located in upland areas near the edge of asettlement system benefited from the movement of game animals from sur-rounding areas, where a combination of very low human population (inferredbased on the absence of residential sites) and high productive potential createdhigher large mammal “source” populations. Although the patterns we see in thisstudy are consistent with such an interpretation, additional research is necessaryto investigate this idea. Isotope analysis of large-mammal teeth (to investigatewhether animals grew up near the sites in whose assemblages they are found orin more distant areas) is one possible route to investigating this further.

In addition to the importance of source–sink dynamics, local populationhistory has an important effect on large mammal relative abundance. This effectis demonstrated to some extent in the Zuni area, but is particularly visible inthe Mesa Verde case. Within <80 years of the establishment of large, permanenthuman settlements, artiodactyl relative abundance is dramatically reduced; inthis study, the reduction in the large mammal index is about 50 %. Laterassemblages show even lower artiodactyl relative abundance as artiodactylnumbers continued to decline. This impact is somewhat shocking in both thespeed with which it takes place and in its severity. However, a growing numberof mathematical modeling efforts in archaeology lend support to such a rapidand dramatic decline (Beaver 2007; Johnson 2006; Schollmeyer 2009, 2011),suggesting that environments have a threshold of human harvest intensity beyond

which large game species cannot be harvested without pushing them into a state ofrapid decline from which recovery is quite difficult. Previous models of humanimpacts on large mammal populations as gradual processes may be in need ofrevision.

Broader Implications

The speed with which local artiodactyl populations are reduced by human activitieswill probably come as no surprise to contemporary biologists; studies in the tropicsindicate substantial impacts to desirable food species often occur within one to twodecades of the establishment of sizeable permanent human settlements (Jerozolimskiand Peres 2003). More important, however, is the observation that despite widespreadlocal depletion of artiodactyls in the areas around prehistoric Southwestern villages,these animals were never locally extirpated. Instead, they remain present in smallnumbers throughout all of the occupations examined here. This may be a highlydesirable outcome in some areas currently facing conservation challenges. Manage-ment strategies often need to allow for continued human hunting; in some moderninstances, bushmeat hunting provides virtually all of the meat in local people’s diets(Peres and Nascimento 2006; Wilkie 2006). However, this often needs to be balancedwith a desire to prevent regional game populations from being pushed to a point ofdepletion from which they cannot recover in order to save some species from localextirpation or even extinction.

Understanding the processes that allowed artiodactyls to persist at a regionalscale in the prehistoric Southwest despite widespread localized depletion mayhelp modern conservation efforts by providing data from a longer time periodthan is possible using contemporary observations. A number of recent conser-vation biology studies identify the protection of game source areas as apotentially promising avenue for protecting dwindling wildlife populations,particularly in areas where enforcing hunting seasons, quotas, and other typesof limits pose many practical difficulties (Fimbel et al. 2000; Hart 2000;Milner-Gulland et al. 2003; Naranjo and Bodmer 2007; Novaro et al. 2005; Peres2001). The majority of these studies have been done in the tropics, an environmentvery different from the case studies considered here. However, the suggestion thatsource–sink dynamics contributed to the long-term resilience of regional artiodactylpopulations in the US Southwest offers some hope that the same process may allowlong-term coexistence of hunters and game populations in other areas. Much workremains to be done on this possibility, and we must be cautious in projecting patternsfrom one area and time period into the present. However, this study highlights thepotential of protecting source areas as one promising avenue for researchers workingon modern protection efforts to explore in the search for long-term solutions.

Acknowledgments We thank Keith Kintigh and Matt Peeples for their assistance with the Zuni case andScott Ortman and the researchers and staff of the Crow Canyon Archaeological Center for their assistancewith the Mesa Verde case. We also thank the journal editors and two anonymous reviewers for theircomments on an earlier draft of this paper. Funding for this research was provided by the Social Science andHumanities Research Council (Canada). Additional support was provided by Simon Fraser University andthe School of Human Evolution and Social Change, Arizona State University.

Casestud

Period(A

Meanelevation

Artiodactyl

blageNISP

References

Buckaroo

Eastern

–1200

Eastern

–1200

RonniePueblo

Eastern

–1200

LizardTerrace

Eastern

–1200

MountainLionHam

Eastern

–1200

Phelps

Eastern

–1200

FlyingFishVillage

Eastern

1000–1130

JuniperVillage

Eastern

1000–1130

AvilasCanyonVillage

Eastern

1000–1130

WellVillage

Eastern

1000–1130

Mitchell

Valley

1000–1130

231,010

JacksonFraction

Valley

1000–1130

Montezuma

Valley

1000–1130

Mattocks

Valley

1000–1130

Valley

1000–1130

Bonito

1300–1

Hinkson

munity

1225–1

Jaralosa

Pueblo

1225–1

Heshotauthla

1325–1

Mirabal

1275–1

BacaPueblo

1300–1

Tinaja

1250–1

RuddCreek

1250–1

Appendix

(contin

Period(A

Meanelevation

Artiodactyl

blageNISP

References

Atsinna

1275–1

Pueblode

losMuertos

1275–1

12,378

RattlesnakePoint

Pueblo

1325–1

39,707

GigantesCom

munity

–1275

Cienega

1275–1

ScribeSCom

munity

–1325

Techado

Spring

–1325

Duckfoot

MesaVerde

840–880

andLeaves

MesaVerde

920–980

16,926

Shields

MesaVerde

1020–1

Shields

MesaVerde

1060–1140

12,913

Shields

MesaVerde

–1225

11,974

Shields

MesaVerde

1225–1

AlbertPorter

MesaVerde

1060–1140

AlbertPorter

MesaVerde

–1225

AlbertPorter

MesaVerde

1225–1

Yellow

Jacket

MesaVerde

1060–1140

Yellow

Jacket

MesaVerde

1225–1

Castle

MesaVerde

1256–1

SandCanyon(secondary

refuse

MesaVerde

1250–1

19,20,21

1Schollm

9);2Schollm

etal.(20

11);3Pow

ell(197

7);4Sanchez

2);5Shaffer

1);6Clark

0b);7Clark

0a);8Thiel(201

0);9Clark

0c);10

7);11Tawater

2Walker(199

etal.(20

11);15

Badenho

rst(20

08);16

Badenho

rsteta

l.(2011);1

7Muireta

l.(2011b);18

DriverandSchollm

(2011);19

Muiret

al.(2011a);20

Muir(199

Kuckelm

an(201

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