PREY BODY SIZE and RANKING in ZOOARCHAEOLOGY: THEORY, EMPIRICAL EVIDENCE, and APPLICATIONS from the NORTHERN GREAT BASIN Author(S): Jack M

Society for American Archaeology

PREY BODY SIZE AND RANKING IN ZOOARCHAEOLOGY: THEORY, EMPIRICAL EVIDENCE, AND APPLICATIONS FROM THE NORTHERN GREAT BASIN Author(s): Jack M. Broughton, Michael D. Cannon, Frank E. Bayham and David A. Byers Source: American Antiquity, Vol. 76, No. 3 (July 2011), pp. 403-428 Published by: Society for American Archaeology Stable URL: http://www.jstor.org/stable/41331900 . Accessed: 26/11/2014 13:19

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This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions PREY BODY SIZE AND RANKING IN ZOOARCHAEOLOGY: THEORY, EMPIRICAL EVIDENCE, AND APPLICATIONS FROM THE NORTHERN GREAT BASIN

JackM. Broughton,Michael D. Cannon,Frank E. Bayham,and David A. Byers

Theuse of body size as anindex of prey rank in zooarchaeology hasfostered a widely applied approach tounderstanding variabilityinforaging efficiency. Thisapproach has, however, been critiqued - most recently bythe suggestion that large preyhave high probabilities offailed pursuits. Here, we clarify the logic and history ofusing body size as a measureofprey rankand summarize empirical data on the body size-return rate relationship. Withfew exceptions, these data document strongpositive relationships between prey size and return rate. We then illustrate, with studies from the Great Basin, the utilityofbody size-based abundance indices (e.g., the Artiodactyl Index ) when used as onecomponent ofmultidimensional analysesofprehistoric dietbreadth. Weuse foraging theory toderive predictions about Holocene variability indiet breadth andtest those predictions using the Artiodactyl Index and over a dozenother archaeological indices. The results indicate closefits between the predictions andthe data and thus support the use of body size -based abundance indices as measures offoraging efficiency. These conclusions have implications forreconstructions ofHolocene trends inlarge game hunting inwestern North America and for zooarchaeological applications offoraging theory ingeneral.

El usodel tamaño del cuerpo como un índice de rango presa en zooarqueológico aplicaciones dela teoríadel forrajeo ha fomentadounenfoque ampliamente aplicado a la variabilidaddela comprensiónenla eficiencia deforrajeo yamplitud dela dieta.Este enfoque hasido criticado periódicamente, conuna preocupación másrecientes derivados delos datos etnográfi- cosque sugieren que la presa de gran tamaño puede estar asociado con altas probabilidades deactividades ypor lo tanto no lastasas de retorno bajo. En este documento, clarificar la lógica y la historiadela utilizacióndetamaño corporal como una medidade rango de presas yresumir losdatos empíricos sobre el tamaño de la relacióncuerpo-la tasa de retorno, haciendo hincapiéenlos artiodáctilos ylos lagomorfos. Estos documento dedatos fuertes relaciones positivas entre el tamañode las presasy tasade retorno, con excepciones limitadas a los casos que no incluyen losartiodáctilos, yque se caracterizan por estrechosmárgenes detamaños delas presas. Acontinuación, ilustran con estudios decaso de la cuencadel norte de la Gran lautilidad del tamaño corporal basado en los índices de abundancia (por ejemplo, elíndice de Artiodáctilo: 2 NISP Artiodác- tilos&[NISPLagomorfos + Artiodáctilos]) cuando se utilizacomo uncomponente deanálisis multidimensionales deampli- tudde la dietaprehistórica. Enestos estudios decaso, utilizamos modelos dela teoríade forrajeo para obtener predicciones sobrela variabilidaddelHoloceno en la amplituddela dietade la hipótesisdeclima tendencias determinadas enla densi- dadde artiodáctilos. Acontinuación, prueba de las predicciones utilizando elíndice de artiodáctilos ymás de una docena de otrosíndices derivados dela fauna arqueológica, floral, yensamblajes dela herramienta. Losresultados indican una estrecha encajaentre las predicciones ylos datos empíricos ycon ello proporcionar unfuerte apoyo para la utilizacióndeltamaño corporalbasado en los índices de abundancia como medidas de eficiencia deforrajeo. Estas conclusiones tienen implica- cionesde largo alcance no sólo para las reconstrucciones delas tendencias recientes delHoloceno enla caza mayor yla vari- abilidaddel comportamiento relacionados enel oeste de América del Norte, pero para las aplicaciones dela teoria del forrajeo zooarqueológicoengeneral.

thepast several decades, models from evo- evolutionin earlyHomo , to theorigins of agri- lutionaryecology have been applied with in- culture,to Mayan monumentconstruction (see In creasingsophistication to a widerange of ar- reviewsin Birdand O'Connell2006; Broughton chaeological issues rangingfrom life-history andCannon 2010). The approachhas becomees- JackM. Broughton ■Department ofAnthropology, University ofUtah, 270 S. 1400E., Room 102, Salt Lake City, Utah 84112 ([email protected]) MichaelD. Cannon■ SWCA Environmental Consultants, SaltLake City, Utah 841 12 FrankE. Bayham■Department ofAnthropology, California State University, Chico, California 85929-0400 DavidA. Byers ■ Department ofSociology, Anthropology andCriminology, Missouri State University, Springfield, Missouri. AmericanAntiquity 76(3), 201 1, pp. 403^28 Copyright©201 1by the Society for American Archaeology

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This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 404 AMERICANANTIQUITY [Vol.76, No. 3, 2011 peciallywell-developed - reaching almost main- Whilethe use ofbody size as a measureof prey streamstatus - in zooarchaeology,the context of rankhas fostered a robust and successful approach severalof the earliest applications of evolutionary to understandingvariation in the archaeofaunal ecology in archaeology(Bayham 1977, 1979, record,most often operationalized through the 1982;Beaton 1973). In thiscontext, foraging the- calculationof body-sized-based taxonomie "abun- orymodels have been effectively applied as tools dance indices" (e.g., Bayham 1979, 1982; forunderstanding prehistoric behavioral variabil- Broughton1994a, 1994b, 2004; Broughtonet al. ityand diachronic trends in foraging efficiency and 2010; Broughtonet al. 2007; Butler2000; Butler dietbreadth. Such variabilityand trendshave, in andCampbell 2004; Byersand Broughton 2004; turn,been linked to many broader developments in Byerset al. 2005; Byersand Smith 2007; Cannon human behavior and morphology,including 2003; Faith2007; Grayson2001; Graysonand changesinvolving technology, the division of la- Delpech1998; Hildebrandt and Jones 2002; Jones bor,the emergence of agriculture,population re- et al. 2008; Kennett2005; Lyman2003a, 2003b; placements,violence and warfare, social inequal- Morin2004, 201 1 ; Nagaoka2002a, 2002b, 2005, ities,settlement systems, material display, and 2006; Porcasiet al. 2000; Ugan 2005a, 2005b; humanhealth (e.g., Bird and O'Connell 2006; Uganand Bright 2001; Wolverton 2005; Wolver- Broughtonand O'Connell 1999; Broughton et al. tonet al. 2008), theapproach has also been cri- 2010; Cannon2003; Fitzhugh2003; Jonesand tiquedperiodically over the past few decades. The Raab2004; Kennett 2005; Kuhn et al. 2001; Morin critiqueshave involved concerns about the effects 2004,2010; Uganet al. 2003). of mass-capturetechniques on preyreturn rates Centralto theseapplications has been theef- (e.g., Madsen and Schmitt1998; Rick and Er- fectiveestimation of keyparameters of foraging landson2000; see also Lupo2007), the high travel theorymodels and articulation of these to theim- costspresumed to be associatedwith large-game perfectlypreserved archaeological residues of past exploitation(McGuire et al. 2007; butsee Grim- humanforaging decisions. Perhaps most important stead 2010), and, most recently, the high mobility in thiscontext is the estimationof preyranks , oflarge game, which is presumedto result in both whichare essentialto applicationof the fine- highpursuit costs and high probabilities of failed grainedprey model, the most widely applied of all pursuits(Bird et al. 2009). Thislast issue, emerg- foragingmodels. Prey-rank estimates are typically ing fromethnographic data collectedfrom the definedin standard model formulations as post-en- Martuof westernAustralia, has led Birdet al. to counterreturn rates, measured as e/h,where e rep- contendthat failure to incorporatepursuit costs resentsthe net energy gain provided by a resource intothe ranking of a preyitem may well compro- andh representsthe handling costs associated with mise"interpretations of variability in prehistoric acquiringand processingit. Of thesevariables, resourceuse" (2009:8). Apparently reflecting these onlyenergy gain is moreor less directlymeasur- concerns,Bird and O'Connell (2006) referto the ablein archaeological contexts because it is directly use of bodysize as an indexof preyrank in ar- proportionaltoprey body size. Because most of the chaeofaunalapplications as a "fragile"assump- handlingcosts are highly variable, context-depen- tion, a conclusionthat is verydifferent from the dent,and difficult to estimatefrom archaeofaunal one we reachhere. materials,prey body size has been commonly used In thispaper, we discussthe role of prey-rank as a proxymeasure of prey rank in the deployment estimatesas one of manyassumptions contained ofthe prey model in archaeofaunalcontexts. This inprey model applications. We clarify the original conventionwas initiallyadvanced in early archae- logicof using body size as a hypotheticalmeasure ologicalapplications of theprey model (Bayham ofprey rank in archaeological applications of for- 1977, 1979, 1982) and has morerecently been agingmodels, and we summarizeavailable em- bolsteredby extensiveethnographic and experi- piricalevidence that speaks to the utility and lim- mentaldata sets bearingon therelationship be- itationsof prey body size whenused in thisway. tweenprey size and post-encounterreturn rate We also explorethe potential influence of mobil- (e.g.,Alvard 1993; Hill et al. 1987;Simms 1985, ity-relatedcosts on thereturn rates for the North 1987;Smith 1991; Winterhaider 1981). Americanmammalian prey most commonly used

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 405 in body size-based abundance indices- cessingtime. Among those, abundance of prey artiodactylsand lagomorphs- and we illustrate inthe diet and size of the prey item are the only withcase studiesfrom the northern Great Basin directquantifiable variables that the faunal therole that abundance indices can play in broader, analysthas access to. The attempt ismade here multidimensionalanalyses of prehistoricforag- tocharacterize the prehistoric diet using size ingbehavior. The mainpoints that we makehere and abundance,thereby facilitating compar- are(1) thatthe relationship between prey body size isonsfrom one siteto another...In thearid andpost-encounter return rate is thoroughlysup- Southwestit is easy to understandthe con- portedfor a widerange of preytypes, especially ceptualutility of this scheme, when we con- thosemost commonly employed in foragingthe- sider the most dominantanimals in the ory-inspiredabundance indices; and (2) thatcon- prehistoricdiet, deer and rabbits. Itis assumed clusionsabout variability in prehistoricforaging thatdeer are the most preferred food item, and efficiencythat are based on patternsobserved in therefore,the representation ofrabbit species abundanceindices are frequentlywell-supported in thediet is an indirectindex of how abun- by other,independent archaeological measures. dantdeer were [Bayham 1977:357]. These have not points far-reachingimplications Furtherwork thatratios of the forrecent reconstructions oftrans-Holocene emphasized only abundancesof such taxa could be extendedto trendsin largegame huntingin westernNorth measuresof hunting efficiency, with high Americaand the debate about whether those trends provide proportionateabundances of large-sized taxa (e.g., aredriven by foraging efficiency or by factors re- deer)indicating higher overall return rates (Bay- lated to and rivalry costly signaling prestige ham 1982, 1986; Szuterand Bayham 1989). Us- (Broughtonand Bayham2003; Broughtonet al. ing measures such as the ArtiodactylIndex 2008; and 2004; Hildebrandt Byers Broughton (2NISP Artiodactyls/SNISP[Artiodactyls + and McGuire2002; McGuireand Hildebrandt Lagomorphs]),Bayham (1977, 1979,1982, 1986; 2005),but also forzooarchaeological applications Szuterand Bayham 1989) explored patterning in of in foragingtheory general. Southwesternhunting efficiency related to a varietyof factors including resource depression and Body Size and PreyRanking in changesin settlementsystem organization. Zooarchaeology:The OriginalRationale Atthe same time, analogous approaches were beingdeveloped in earlyapplications of the prey ofthe of size Recognition importance preybody modelwithin the field of zoology. Wilson (1976), as a variableinfluencing food choice significant forinstance, argued that the abundance of large, has a in White longhistory zooarchaeology(e.g., relativeto small,insect prey found in the stomach even to itsuse as a for 1952, 1953), prior proxy contentsof birdscould be used to measurethe rankin early foraging theory applications in prey qualityof theirforaging environments and thus thefield. As a characteristicthat is readilyde- prey foragingefficiency. Since large, high-ranked prey rivedfrom faunas,the use ofbody archaeological shouldalways be pursued,Wilson (1976:96) ar- size in thiscontext outof the limitations of grew gued,their relative abundances measure both how earlierbiomass-based efforts to understand the se- frequentlythey were encountered and the amount lectiveutilization of animals, which data required ofenergy "available in theenvironment." difficultto derivefrom settings archaeological We emphasizethat these structurally similar 1974; Munsonet al. 1971; Smith (e.g., Grayson earlyprey model applications in zooarchaeology 1974,1979; Yesner and 1976;see Bayham Aigner and zoologyboth fully recognized the potential 1982 forfurther discussion). Pioneering the ar- importanceof pursuit costs, but, since they dealt ticulationof model witharchae- prey predictions onlywith the residues of past foraging decisions, measuresof the relative abundance of dif- ological neitherhad any secure way to accessthem. Thus, ferent-sized reasoned: preyspecies, Bayham adoptinga standardmodeling convention, vari- - It is clearthat a varietyof factorsinfluence ables perceivedto be intractablealbeit poten- whichanimals are selected for food, such as tiallysignificant - were held constantto allow size,density, palatability, search time and pro- theanalysis to proceed(see also Griffiths1975).

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 406 AMERICANANTIQUITY [Vol.76, No. 3, 2011 Bothapproaches therefore adopted body size as a To ourknowledge, there are now ten data sets proxyfor prey rank given its intrinsic link with en- in existencethat include return-rate estimates for ergeticgain. suitesof vertebrateprey types consisting of five We also emphasizethat the assumptions about or moretaxa or preytypes; these data sets are preyranks incorporated in thistradition of prey- summarizedin Table 1. In all of thesecases, the model applicationsin zooarchaeologyhave al- directionof the relationshipbetween post-en- waysbeen viewed as hypothesesthat enable the counterreturn rate and body size is positive,and generationof testablepredictions, rather than as in eightof the ten cases thecorrelation is signif- absoluteor deterministic characterizations ofpast icantat an alphalevel of .10 (Table 1). Because humanforaging behavior. As withall modelcom- theeffects of prey mobility and failed pursuits on ponents(currency, constraints, etc.), the assump- returnrates have been recently highlighted (Bird tionsmade about prey rankings are at riskwhen- et al. 2009), we pointout thatall of theethno- evera modelis appliedin a particularcontext. graphic-observationaldata sets representedin Predictivefailures imply that one or moreof the Table 1 explicitlyinclude the costs of failed pur- assumptionsincorporated into a model,including suitsin theirreturn rate estimates. Since signifi- thoseassociated with prey rankings, may be in- cantcorrelations between prey body size andre- appropriate.Predictive successes, on the other turnrates are indicatedin almostof all of these hand,imply that model assumptionsare valid. cases,it would appear that prey mobility may be Thewide range of successful tests involving mod- less influentialthan has beensuggested. els thatassume maximization of foragingeffi- Pursuingthis further, the two empirical cases ciencyas a goal and currency,and that use body thatfail to producesignificant correlations be- size as a proxyfor prey ranks, attest to the gener- tweenprey body size and returnrate - thosein- alityof these particular assumptions. volvingthe Mayangna-Miskito and Martu- are instructivein thatthey elucidate an important factorthat to Ethnographicand ExperimentalEvidence general appears affectthe body size-returnrate Supportthe Body Size-ReturnRate relationship.Specifically, com- todata sets derived from other Relationship pared casesthat in- volve terrestrialhunting, these two incorporate The firstwave of preymodel applicationsin verynarrow prey size rangesand very small max- zooarchaeologyreasoned that, though context- imumprey sizes: the Martu data, for example, ex- dependent,pursuit costs would typically not vary hibitthe lowestrange of exploitedprey sizes, to such a degreethat an ordinalscale ranking with hill kangaroo(Macropus robustus),the based on size alone wouldbecome reversed, es- largestprey type, weighing only 22 kgand cossid peciallyfor taxa of suchdisparate size as artio- larva,the smallesttype, weighing just 13 g. In dactylsand lagomorphs.This conclusionbegan contrast,the prey-size range for the Cree is over to be empiricallysupported as soon as ethno- 415 kg, withmoose ( Alces alces) and grouse graphicand experimentalanalyses of post-en- (Phasianidae),representing the largest and small- counterreturn rates startedto emergein the est preysizes, respectively(Table 1). We also 1980s(e.g., Hawkes et al. 1982; Hill et al. 1987; notethat the Mayangna-Miskito and Martu cases Simms1985, 1987; Winterhaider1981). By the are theonly two involving terrestrial hunting in early 1990s, summariesof the empiricaldata whichartiodactyls are not included in the suite of on therelationship between vertebrate prey body preyexploited. Thus, we can onlyconclude that size andreturn rate showed positive correlations thelack oftruly large-bodied prey and therela- in each and everycase (e.g., Broughton1994a). tivelynarrow range of preysizes in thesetwo These data suppliedestimates for other critical cases mustaccentuate the effect of variationin model parametersand enabled a surgein the preymobility on prey rankings. This, of course, is applicationof foraging models not only in zooar- nota new point,having previously been articu- but chaeology throughoutarchaeology in general latedmost notably by Stiner and colleagues (e.g., (see Bird and O'Connell 2006; Cannon and Stineret al. 1999; Stineret al. 2000; Stinerand Broughton2010). Munro2002; see also Morin2011), who have

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This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 408 AMERICANANTIQUITY [Vol.76, No. 3, 2011

Americanarchaeological applications of foraging theory.We thenshow how concernsabout the generalityof the body size-return rate relationship can be furtheralleviated by employing body-size- basedabundance indices as onlyone ofa battery ofarchaeological and paleoenvironmental indices in testsof hypothesesabout foraging efficiency.

Mobility,Return Rates, and North AmericanTerrestrial Mammals: Artiodactylsand Lagomorphs

Figure1. Relationshipbetween log range of prey weight Artiodactylsand lagomorphs were primary faunal withina dataset and thestrength of thecorrelation resources used by prehistoricpeoples in North betweenbody size and return rate (from terrestrial prey and dominate if datasets in Table 1). America, collectivelythey many, notmost, archaeological faunas from the interior of thecontinent. These preytypes differ consid- pointedout thatpost-encounter return rates for erablyin body mass and werethe focus of the small-bodiedprey might vary considerably due to originalforaging theory-based abundance index, variationin preymobility. Indeed, in theentire theArtiodactyl Index, which has since become the sampleof studies that include terrestrial hunting, single-mostwidely applied abundance index used thesize rangeof exploited prey is positivelycor- totrack foraging efficiency and diet breadth. The relatedwith the strengthof the correlationob- ArtiodactylIndex has figuredprominently in re- servedbetween body size andpost-encounter re- centreconstructions of trans-Holocene trends in turnrate ( rs= .73,p = .04; Figure1; see Morin large-gamehunting in westernNorth America 2011 forfurther discussion). That is, thestudies and the associateddebate about whether those thatincorporate a greater range of prey sizes tend trendsare drivensolely by concernsof foraging to producestronger correlations between body efficiencyor, alternatively, by factorsrelated to size andreturn rate, suggesting that variability in matingeffort, costly signaling, and prestige rivalry factorssuch as preymobility is likelyto con- (Broughtonand Bayham 2003; Broughtonet al. foundthe general body size-prey rank relation- 2008; Byersand Broughton2004; Hildebrandt shiponly when the range of bodysizes involved and McGuire2002; McGuireand Hildebrandt is smallto beginwith. 2005).While much of this debate has centered on In sum,the logic outlined above and the com- theeconomics associated with harvesting artio- plementaryempirical support just summarized dactylsand lagomorphs, the potential influence of providea securefoundation for using body size as differencesin mobilitybetween artiodactyls and a proxymeasure of preyrank in manycases. In lagomorphson theirreturn rates has notbeen ex- turn,this provides a basis forconstructing and aminedin detail. testingspecific hypotheses about variabilityin Failed overallforaging efficiency, as measuredby the Mobility; Pursuits, and ReturnRates relativeabundances of different-sized prey, in re- forArtiodactyls and Lagomorphs lationto trendsin humandemography and/or en- To beginsuch an examination,we firstpoint out vironmentalchange. To be sure,the empirical that,using the classificationof Bird et al. data do suggestthat attention should be paidto the (2009:11), all artiodactyland lagomorphtaxa magnitudeof therange in bodysize amongthe wouldbe considered"fast" prey - thehighest cat- preytypes that are considered in any analysis. We egoryon theirscale of preymobility. The maxi- next illustratehow thismight be takeninto ac- mumrunning speeds for black-tailed jackrabbits count by exploringin furtherdetail the relevant (Lepus californicus), for example, exceed 64 characteristicsof artiodactylsand lagomorphs, km/hr,falling within the range of speeds for most thevertebrate taxa most often considered in North artiodactyls,including mule deer ( Odocoileus

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 409 studies,data on thenumber of animalspursued per hunt- that is, animals eithershot at or "jumped"- are presentedalong withthe total numberof animals harvested, enabling the probabilityof successful pursuits to be estimated.The dataon lagomorphsinclude 1 1- and 3-year survey periodsfor cottontail (Sylvilagus spp.) hunters fromSouth Carolina and Kentucky,respectively. The artiodactyldata are forwhite-tailed deer ( Odocoileus virginianus)hunters from single- yearsurveys in Ontarioand SouthCarolina. Theaverage pursuit success rate for 14 years of Figure2. Relationshipbetween log weight and return rate surveydata for rabbits is .56, withyearly values forartiodactyls andlagomorphs. Data from references in thatrange between .62 and .42. This meansthat Table1 and Ugan(2005a). lagomorphsescape from modern shotgun hunters between60 percentto 40 percentof the time. The twodifferent white-tailed deer studies produced hemionus),elk ( Cervuscanadensis ), bison{Bison nearlyidentical pursuit success rates of .82 and bison),and bighorn sheep (Ovis canadensis) (Gar- .79,with a meanof .80. Failedpursuits thus occur land1983; Garland and Janis 1993; Morin 201 1). in onlyabout 20 percentof encounters.The sub- The greatagility and elusiveness of lagomorphs, stantialdifferences in targetsize and thegreater althoughdifficult to quantify,is also legendary visibilityof artiodactyls within areas of dense veg- andmay be enhancedby their smaller size relative etation,we suspect,may lie atthe heart of the dif- toartiodactyls. These considerations suggest that ferencein pursuitsuccess betweenartiodactyls pursuitcosts and the probability of failed pursuits and lagomorphs.In anycase, thesedata are con- mightbe broadlycomparable for artiodactyls and sistentwith the ethnographically derived return- lagomorphs- or even higher for lagomorphs - all rateestimates in thatthey indicate no overlapin else beingequal. If so, thenpost-encounter return pursuitsuccess rates between rabbits and deer. ratesfor artiodactyls and lagomorphswould, on We are underno illusionthat these actualisti- average,be largelya functionof thesubstantial callyderived return-rate and pursuit-success data differencesin bodysize thatseparate the two or- are necessarilyrepresentative of the actual ab- ders.This is, in fact, born out by the available em- solutevalues associated with prehistoric foragers piricaldata on returnsfor these taxa. in anyparticular context. They do, however,pro- Figure2 showsthe relationship between log videno challengeto using prey size, the most im- prey-bodysize and empiricallyderived post-en- portantknowable economic quality of artiodactyls counterreturn rates for the entire sample of North andlagomorphs, as a sourceto hypothesize an or- Americanartiodactyls and lagomorphs. These in- dinal-scaleranking of them in archaeofaunal stud- cludeall of therelevant data from the studies in ies. Ifanything, they suggest that lagomorphs are Table1 as wellas severalreturn-rate estimates for generallymore difficult to capturethan artio- lagomorphsprovided in Ugan(2005a). The rela- dactyls,which only reinforces the point that lago- tionshipis positiveand highly significant (r2 = .80, morphsshould, in general, provide lower post-en- p < .0001; rs= .84,p = .0004),with dramatically counterreturn rates than artiodactyls. significantdifferences in meanreturn rates be- Modelingthe Effect of Pursuit tweenthe two orders ( t = 6.58,df= 17, p < .0001 ). Successon ReturnRates Furtherinformation onmobility-related pursuit costsare available from studies involving modern This analysiscan be madesomewhat more gen- the lagomorphand artiodactylhunters in eastern eralby deconstructing, on a theoreticallevel, NorthAmerica (Hols worth 1973; Morgan2005; effectof preycapture success on post-encounter Ruthand Simmons1999; South CarolinaDe- returnrates. As notedabove, in standardformu- arecalculated partmentof NaturalResources 2009). In these lations,post-encounter return rates

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 410 AMERICANANTIQUITY [Vol.76, No. 3, 2011 as elh,or energeticgain per handling time. As a greaterfor lagomorphs than for artiodactyls for characteristicof a preytype, the quantity elh is the averagepost-encounter return rate of lago- generallyrecognized to be an expectedor average morphsto exceedthat of artiodactyls.Given the value(e.g., Stephensand Krebs1986:13-17), in modernhunting survey data discussedabove, lightof the fact that there may be someintra-prey- which suggest that pursuit success is likelyto be typevariability inreturn rate, due to, among other lowerfor lagomorphs than for artiodactyls (per- factors,success or failure in pursuit. The effectof hapsby half), it is difficultto see howprehistoric capturesuccess on preyranks can be understood lagomorphcapture success could everhave ex- moreexplicitly by substituting into the calculation ceededthat for artiodactyls by sucha margin. ofthe post-encounter return rate the equality e = where is the of /?*g, p probability successfully MultipleQuantitative Indices ofDiet a and is the ener- capturing preytype g average Breadthand ForagingEfficiency geticgain provided by that prey type when pursuit is successful.Thus, for prey type /, the average The data just discussed stronglysuggest that post-encounterreturn rate (r) canbe calculatedas analysesof patternsin foragingefficiency that ri- (Pi*gi)/hi9with h representinghandling time relyon thebody size-return rate relationship, at averagedacross both successful and unsuccess- leastas appliedto artiodactylsand lagomorphs, fulpursuits.1 are valid. However,as in any archaeological To illustrate,a prey type that could be captured analysis,even stronger cases regardingtrends in withoutfail (i.e., p- 1) withan averageenergetic foragingefficiency can be madewhen multiple, valueupon capture (g ) of 1,000kcal and an aver- independentlines of evidenceare employed.In age handlingtime of 1 hrwould provide an aver- otherwords, a singleabundance index is onlyone age post-encounterreturn rate of 1,000kcal/hr. of manyindividually less-than-fool-proof tools The exact same averagepost-encounter return thatcan be andhave been used in thesekinds of ratewould also be providedby a larger-bodiedbut analyses.It is thereforeworth noting that the moreelusive prey type with an averageenergetic mostcompelling examples of prehistoric changes gainof 10,000kcal and an averagehandling time in foragingefficiency do notrely solely on the of 1 hrthat could only be capturedin one outof body size-returnrate relationship, but instead tenpursuits (p = .1). Whenthe effects of capture havebeen demonstrated through the use ofmul- successare made explicit in this manner, it can be tiplemeasures that collectively signal compara- seenthat the average post-encounter return rate of ble trends.Zooarchaeological complements to a large-bodiedprey type might indeed be lower bodysize-based abundance indices may include, thanthat of a smaller-bodiedprey type if the butare notlimited to, measures of evennessand probabilityof capture success is sufficientlylower richnessas indicatorsof faunaldiet breadth; forthe former than for the latter. Specifically, if skeletalpart representation reflecting local de- preytype 1 is thelarge-bodied type and prey type pressionand distant patch use; age, size,and sex 2 is thesmall-bodied type, then for prey type 2 to structureas theyrelate to harvestpressure, prey providethe higher average post-encounter return depression,and habitat quality (e.g., Stiner et al. rate,it must be thecase that 2000; Wolvertonet al. 2008); and bone frag- mentationand othervariables that may relate to ÜL)f + processingintensity (Ugan 2005b). Abundance Pi Ui fh/ indicesconstructed to reflectthe harvesting of preytypes from proximal and distantpatches Applyingthis to artiodactylsand lagomorphs, have also playeda keyrole in documentingde- the dataused to generate Figure 2 suggestaverage cliningreturns for central-place foragers in a values of g and /г,respectively, on theorder of numberof studies(Broughton 1999, 2002; Na- 51,000 kcal and 2.35 hoursfor artiodactyls and gaoka 2002b). Indicesconstructed to represent 1200kcal and 6 minutesfor lagomorphs. In turn, therelative frequency of highand low ranking thiswould suggestthat the probabilityof suc- small-sizedprey have also been effectivelyde- cessfulcapture would have to be atleast 1.8 times ployed,in manycases relyingon distinctivedif-

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 411 ferencesin preymobility (Stiner et al. 2000; dices used to independentlyinform on foraging Stutzet al. 2009; Wolverton2005). efficiencyand prey abundances in thesesettings. In additionto thesevariables - now industry We nextfurther illustrate the strength of the "mul- standardsin zooarchaeologicalapplications of tiplelines of evidenceapproach" with recent re- foragingtheory - trendsin frequenciesof tools searchand ongoing analyses involving Holocene (toolabundance indices) associated with the har- climatechange and trends in human hunting effi- vestingof particular resources can be established ciencyin thenorthern Great Basin. to furthertest hypothesized trends in preyabun- dances.In more exceptionalcases, independent Holocene ClimateChange and Human Diet datacan also paleontological providesupporting Breadthand ForagingEfficiency in the evidence,as inthe example we discussbelow in- NorthernGreat Basin volvingthe trans-Holocene artiodactyl fecal pelletrecord from Homestead Cave, Utah.Quanti- In recentwork, we testedthe hypothesis that the tativepaleoclimatic indices of manykinds have seasonalityof temperatureand precipitation also longbeen employed in the context of tests in- playeda majorrole in controllingthe population volvingtrends in foragingreturns. And finally, densitiesof artiodactyls (e.g., bighorn sheep [Ovis trendsin humanskeletal paleopathologies and canadensis], muledeer [Odocoileus hemionus], staturehave also beenlinked to trends in foraging and pronghorn[Antilocapra americana]) across efficiencyderived from faunal indices: insofar as theterminal Pleistocene and Holocene of western lowerforaging efficiency implies greater foraging NorthAmerica (Broughton et al. 2008). Formuch effortrequired to meetminimum caloric require- ofthis region, general circulation climate models mentsand an increasedrisk of malnutrition,and a rangeof paleoclimatic data suggest that sea- higherlevels of morbidityand mortality,slower sonalextremes in temperaturepeaked during the growthrates, and reduced adult body size should terminalPleistocene and early Holocene and that accompanydeclines in the former(Bartelink earlyand middle Holocene precipitation followed - 2006;Broughton and O'Connell 1999; Broughton a winter-wet,summer-dry pattern conditions et al. 2010). knownto depressartiodactyl densities. These To thesemore established measures we can trendsare mirroredin a macrophysicalclimate addpromising but still experimental use offaunal simulationmodel (MCM) developedfor the north- boneisotopie and ancient mtDNA analyses. Bone ern BonnevilleBasin (Figure3), northwestern isotopeanalyses have been used to determinethe Utah,from which we derivedterminal Pleistocene geographicorigin of preyanimals and identify andHolocene climatic values and three indices of possibleinstances of distant patch utilization dri- climaticseasonality: (1) intra-annualtemperature venby local patchdepression (Grimstead 2005, range,(2) summerprecipitation intensity, and (3) 2009). AncientmtDNA analyses of faunalbone winterprecipitation intensity. These indices were can potentiallyreveal trends in geneticdiversity thenarrayed against detailed late Quaternary ar- thatmay reflect changes in prey population sizes, tiodactylabundance records in the Bonneville and preliminarywork with California tule elk Basin. These includeda uniquepaleontological and Northernfur seal showspromising signs for recordof fecalpellet densities from Homestead theapproach (Beck 2010). Cave and archaeologicalrecords of artiodactyl Withthe exceptionof these last measures, faunalremains (i.e., an ArtiodactylIndex) and wherework is stillin itsinfancy, good use of inlarge-gamehunting tools from Hogup Cave. Each dices constructedfrom all the othervariables of theseartiodactyl abundance records showed listedabove has beenmade in analysesof trends significantcorrelations with the model-derived in prehistoricforaging behavior the world over, seasonalityindices and suggested that artiodactyls withmost also attentiveto taphonomic issues that occurredin low densitiesfrom the terminal Pleis- whilesub- maycloud their meaning. Table 2 presentsa non- tocenethrough the middle Holocene, sum- comprehensivesampling of studiesin thistradi- stantialincreases occurred during equable, tionthat use body-sizebased abundanceindices, mer-wetperiods of thelate Holocene.Although is alongwith the numbers and types of additional in- geographic,site- and species-specific variability

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This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 413

Figure3. Mapof the northeastern Great Basin showing locations ofsites discussed inthe text. anticipatedand clearly exists (see detaileddiscus- HogupCave : PreviousResearch sionin Broughtonet al. 2008:1931-1932;Byers 2 The deep,well-stratified deposits of Hogup Cave andBroughton 2004; see also Hockett2005), ar- havebeen the focus of renewed interest in the ef- vertebraterecords from across west- chaeological fectof climate change on trans-Holocenehunting ernNorth America showed similar very temporal patternsin the Bonneville Basin (Broughton et al. in that patterns artiodactylabundances, suggesting 2008; Byersand Broughton 2004; Byersand Hill thetrend and itsclimate-based correlates be may 2009; Hockett2005; Figure3). HogupCave is a a one. verygeneral limestonecavern located along the southern end Insofaras artiodactylsrepresented high-return of the Hogup Mountains.Sixteen stratigraphie to ancienthunters in these preytypes settings, unitswere encountered during the excavation of these trendsshould have caused substantial theHogup Cave sediments,which reached over in and dietbreadth. changes huntingefficiency 4 m in depth(Aikens 1970). One-quarter-inch More itcan be thatthe late specifically, predicted screenswere used to collectnot only an enor- Holocene in densitiesshould be spikes artiodactyl moussample of artiodactyl bones and other fau- associatedwith overall efficien- higher hunting nal remains,but also an extensiverecord of per- and theseshould be reflectedin cies, multiple ishable artifacts,including textiles, nets, and additionallines of evidence.In this we section, moccasins(Aikens 1970; Durrant1970; Par- research on Holocene synthesizeprevious bearing malee 1970). Thirty-two,4C dates place thehu- trendsin BonnevilleBasin foragingefficiency manoccupation of the cave betweenabout 8,800 withnew tests to the Cave verte- applied Hogup and 480 yearsB.P.3 Although several of these bratefauna and withnew data fromthe Little dates are out of stratigraphieorder, as others Boulder locatedto thewest of theBon- Basin, have noted(e.g., Grayson1993; Madsen and nevilleBasin in northernNevada. Berry1975; Mullen 1997), the Hogup Cave dates

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Figure5. Comparisonbetween Hogup Cave early/middle andlate Holocene aggregate pronghorn ageprofiles (from Byersand Hill 2009: Table 9).

etal. 2003),which provides an innovativeframe- workfor and subsistence Figure4. Thedistribution ofthe artiodactyl andprojectile linkingtechnological pointindices (standardized) across the Hogup Cave strata. change.This modeldoes so by showingthat the greaterthe amountof timespent harvesting a overallare highlycorrelated with stratigraphie particular resource, the higher the payoff for time position( rs= .77,p < .0001). investedin technology associated with handling it. As previouslydocumented, Artiodactyl Index The modelthus predicts that relative abundances values derivedfrom the Hogup Cave archaeo- of differentkinds of technologiesshould vary in faunalassemblage are consistently low during the tandemwith the relative abundances of thefood earlyand middleHolocene but increase,with resourceswith which they are associated. Figure two markedspikes, during the late Holocene. 4 showsboth the artiodactyl and projectile point This patternis verysimilar to the trenddocu- indicesplotted together across the Hogup Cave mentedin artiodactylpellet densities at Home- strata.The twovariables appear well aligned, and steadCave (Madsen 2000). Not onlyare artio- a correlationanalysis confirms this impression (rs = dactylbones proportionatelymore abundant in .80,p < .01). Further,this tool-based index of the collectiveset of late Holocene stratacom- artiodactylencounter rates and foraging efficiency paredto thoseof the early and middle Holocene, is also correlatedsignificantly with climatic sea- theirfluctuations within the late Holocene are sonalityindices derived from the local climate well alignedwith the artiodactyl pellet spikes at model(Broughton et al. 2008). HomesteadCave. Bothdata setsshow dramatic Recentfine-scale analyses of pronghornde- peaks in artiodactylspecimens between about mographicstructure at Hogup provideno evi- 4,000and 3,000 yrs B.P. andagain between 1,200 dencefor any change that might suggest that these and 1,000B.P. Bothare also significantlycorre- patternsin theabundances of faunalremains or latedwith model-derived indices of climatic sea- toolsare related to a shiftin the functional use of sonality(Broughton et al. 2008). thecave or a changein the dominant mode of pro- These patternsare also consistentwith tem- curement(Byers and Hill 2009; butsee Aikens poraltrends across the Hogup Cave stratain the 1970). Dentition-derivedmortality profiles are relativeabundances of artifacts likely used to har- statisticallyindistinguishable between early/mid- vestartiodactyls and lagomorphs. Insofar as nets dle andlate Holocene assemblages, with both be- andsnares were commonly used to capture lago- ingcharacterized by an attritionalprofile sugges- morphsand projectile points were used to kill ar- tiveof encounterhunting (Figure 5; Byersand tiodactyls,then the projectile point index (2 Pro- Hill 2009). Plannedanalyses of skeletal part rep- jectilePoints/[2 Projectile Points + 2 Cordage]) resentationwill allow furtherevaluation of po- providesyet another measure of artiodactylden- tentialchanges in sitefunction. sitiesand overall hunting efficiency. A theoretical Finally,we note thatprevious taphonomic basisfor the use of suchan indexis givenby the analysesof the Hogup lagomorphassemblage "techinvestment model" (Bright et al. 2002;Ugan provideno suggestionthat these trends are related

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 415 tochanges in the proportionate use ofthe cave by Moreover,the SEI is significantlycorrelated with people and otheragents. Most notably,no sig- theArtiodactyl Index ( rs= .74,/7 < .01), thepro- nificantdifferences are evidentin the relative jectile pointindex ( rs = .79, p < .01), and the frequenciesof raptor/carnivore marks compared threeindices of climaticseasonality (winter pre- to human-induceddamage (cut marks, burning) cipitation:rs = -.73, p < .05; temperaturerange: betweenthe early/middle and late Holoceneas- rs= -.86, p < .01; summerprecipitation: rs= .73, semblages(Byers and Broughton2004; Hockett p < .05),which, again, are hypothesized to be dri- 1993, 1994). vingthe trends in artiodactyldensities and overall huntingefficiency. Finally, we notethat the SEI Cave: NewTests Hogup is uncorrectedwith the underlying strata sample To furthertest whether the trends previously doc- sizes in theHogup collection ( rs = -.34, p > .15), umentedat HogupCave trulyreflect changes in alleviatingany concernsthat might arise were huntingefficiency and diet breadth, we applysev- sucha correlationpresent (e.g., Grayson1984). eral additionalindices to the vertebratefauna Taxonomiediversity and richnessmeasures fromthis site here. These includea selectiveef- can provideadditional indices of diet breadth and ficiencyindex that incorporates the entire assem- foragingefficiency in archaeologicalfaunas, es- blageof identified mammals, rather than only ar- peciallyif used alongside other measures that di- tiodactylsand lagomorphs,and taxonomie rectlyincorporate prey rank estimates. All elsebe- richnessand diversity indices applied to both the ingequal, faunas deposited by foragers with wide mammaland aviancollections from the site (Ta- dietbreadths and low foraging efficiencies should bles 3 and4). be representedby manyprey types, with more Althoughthe trend in theArtiodactyl Index at equal distributionsof abundancesacross them HogupCave is correlatedin theexpected direc- (Grayson 1991; Graysonand Delpech 1998; tionswith indices of seasonality,a tool-based Jones 2004; Nagaoka 2001; but see also measureof foragingefficiency, and the Home- Broughtonand Grayson1993). Conversely,in steadCave pelletrecord, it incorporates only the contextscharacterized by highoverall returns, identifiedartiodactyl and lagomorphspecimens huntersshould focus disproportionately on only a fromthe collection, and these taxa comprise only fewof the highest ranked prey. Thus, as has been 64 percentof the total mammalian MNI fromthe articulatedelsewhere, wide diet breadths should site. More taxonomicallycomprehensive tests be associated with high numbers of taxa thusseem warranted, and one suchapproach is to (NTAXA) andhigh diversity values as measured use a versionof Bayham's(1982, 2010; Szuter by measuressuch as theShannon- Wiener index and Bayham 1989) selectiveefficiency index (H' = -2 Pi logpt). We can thereforehypothesize (SEI). As adaptedhere, the SEI maybe definedas: thatthe early/middleHolocene assemblagesat Hogupshould be characterizedby high NTAXA SE l = (ZMiEi)/ZMi andShannon-Wiener diversity values, while those whereMř is an estimateof the relative abundance forthe late Holocene samples should be charac- ofprey type i andEt is an estimateof theweight terizedby low valuesof these measures. in kg perindividual of preytype i. In additionto Since NTAXA is widelyknown to be highly artiodactylsand lagomorphs, values of this index dependenton samplesize, measured either as to- thusinclude the contributionsof the variably talMNI ortotal NISP (e.g.,Grayson 1984, 1991), abundantrodents and carnivoresin thesequence samplesize mustbe takeninto account when as- and providethe aggregate mean weight of indi- sessingvariation in richness across the Holocene vidual animalsrecovered from any givenstra- depositsat Hogup Cave. The approachthat we tum.Insofar as bodysize is an appropriatemea- takehere to doingthis is to compareregressions assem- sureof prey rank in this setting, this index should of NTAXA on sample size forsets of be low duringthe earlyand middleHolocene blagesthat are hypothesized to differin richness strataand increasein tandemwith other mea- (i.e., to testfor differences in richnessthrough thatare suresduring equable climatic periods of thelate analysisof covariance).Assemblages exhibit Holocene.Figure 6 showsthat it does just this. samplingbroader underlying diets should

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Moisture Winter Summer TemperatureArtiodactyl Index Proj. Pt. Index Stratum Index3 Precip. Precip. Range (Z-score) (Z-score) 1 -40.63 25.80 3.28 31.87 -.856 -.381 2 ...... 856 -1.003 3 -38.08 24.68 3.79 32.46 -.517 -.692 4 -38.27 25.05 3.25 32.20 -.743 -1.003 5 -40.69 25.75 3.11 32.11 -.856 -.817 6 -40.77 25.84 3.40 31.50 -.856 -1.065 7 -40.77 25.84 3.40 31.50 -.856 -.879 8 -40.39 25.69 4.66 30.32 -.517 -.257 9 .... .050 .117 10 -40.11 23.49 8.04 29.88 1.861 1.236 11 - - - 2.088 -.257 12 -39.51 24.09 9.42 29.32 -.177 .925 13 - - - - .729 .552 14 -40.00 22.50 9.28 28.90 1.295 2.294 15 - - - - -.177 -.070 _16 -38.95 22.57 830 28.73 389 1.299 aTheclimatic values presented here are from Broughton etal. (2008). As discussed therein, values were not calculated for stratathat lacked dates (11, 13, 15) or were represented onlyby anomalous ones (2, 9). - higherregression slopes and/orintercepts, indi- densitieswere low butthe differences are not catingthat they contain more taxa, on average,at statisticallysignificant at an alpha level of .05 any given sample size (e.g., Cannon 2004; (Table5). Wedo note,however, that the difference Graysonand Delpech 1998). Figures 7 and8 dis- in elevationfor mammals (p < .08) and thedif- play the relationshipsbetween logarithmically ference in slopefor birds (p = .1 1) are nearlyso. transformedsample size valuesand NTAXA for Turningto diversity, as Figure9 shows,mam- theHogup mammals and birds, respectively, with malianand avian Shannon-Wiener diversity in- regressionlines plotted separately for the dicesfor the Hogup Cave faunaeach decline dra- early/middleand late Holocene assemblages (see maticallyin thelate Holocene from consistently Grayson1991 forthe protocol we used to count higher values during the early and middle "overlapping"taxa). These data suggestgreater Holocene.In fact,comparisons between the col- richnessamong the early/middleHolocene as- lectivesets of early/middleand late Holocene - semblagesthan the late Holocene assemblages strataat Hogup showsignificant differences, in andhence broader diets at times when artiodactyl theexpected directions, in average diversity index

Figure7. Relationshipsbetween NTAXA and sample size Figure6. Distributionof the selective efficiency index forearly/middle andlate Holocene mammalian assem- (SEI)across the Hogup Cave strata. blagesfrom Hogup Cave.

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 418 AMERICANANTIQUITY [Vol.76, No. 3, 2011 14-| 7 For theHogup Cave birdcollection, the de- O Early/MiddleHolocene 77 " Early/MiddleHolocene „ л / clinein thediversity index is potentiallyclouded 12 • LateHolocene // LateHolocene О / by a correlationwith sample size ( rs-.87, p < - О 10 // .001). However,we notethat the pattern of posi- <л / tivecorrelations between size andboth di- I 8- sample versityand richness in theavian data set may ac- Z /О tuallyprovide further insight into an underlying 4" / commoncause of the correlations in thisinstance. It is readilyapparent that bird-bone frequencies declinesubstantially in thelate Holocenestrata оJ , , , , . -0.25000 025 0.50075 1.001 25 150 175 200 (Table3), a periodwhen other indicators suggest LogNISP (Birds) thatforaging efficiency was increasing.Although analysesof the Hogup Figure8. Relationshipsbetween NTAXA and sample size systematictaphonomic forearly/middle and lateHolocene avian assemblages avian assemblage have yet to be conducted,pre- fromHogup Cave liminaryexamination suggests that, while rap- torsundoubtedly made a contribution,many bird bonesexhibit cutmarks or evidence of in- Table5. Test Statistics Comparing thelog Sample Size- burning NTAXARelationships Between the Early/Middle andLate dicativeof human involvement intheir deposition. HoloceneAssemblages from Hogup Cave. Insofaras peopleplayed a substantialrole in the depositionof the Hogup avian fauna, the intensity Mammals Birds ofbird harvesting - relative to, say, large mammal t P t p - could a indexof for- Differencein .06 .47 1.30 .11 hunting provide negative slope boththe small size and Differenceinelevation 1.47 .08 .99 .17 agingefficiency, given Note:All tests are one-tailed. highmobility of thebest represented taxa in the collection(e.g., ducks, eared grebes). Indeed,the total bird NISP perstratum is neg- 3.0 - • - Mammal ativelycorrelated with the Artiodactyl Index (rs = --9- Bird = X 2.5- © -.72, p < .01), themammalian SEI ( rs -.57,p < -о л M jk .05), andthe projectile point index ( rs= -.62,p < Í & ' /' ^ /' ¡' . 2-°- .05). The precipitouslate Holocene drop in bird- *V 'V 1 '/ V j/' bone depositionmay thushave its rootsin the diet-breadthchanges indicated in other aspects of thefauna but at thesame time may be generating the depressionof both NTAXA and diversity Й 0-5■ 1 ' acrossthis period. '' °°-l I HogupCave : Summary ^ V «JQ> «V4- ,$>^чО s> Sevendistinct indices of foragingefficiency and Stratum(Date14C yr.bp.) dietbreadth have now been applied to the Hogup Cave vertebratefauna: two abundanceindices Figure9. Distributionofthe Shannon- Wiener diversity based on preybody size (theArtiodactyl Index indexfor birds and mammals across the Hogup Cave strata. and theSEI), one abundanceindex derived from thehunting tool assemblage,and fourmeasures involvingtaxonomie richness and diversity (two values (mammals:t = 2.13, p = .05; birds:t = each formammals and birds).With only a few 11.87, p <.001). We also observethat there is no ambiguitiesstemming from sample size issues, significantcorrelation between the Shannon- theindices consistently reflect wide diet breadths Wienerindex and samplesize forthe large col- andlow foraging efficiencies throughout the early lectionof mammalmaterials from the site (rs = andmiddle Holocene, with dramatic increases in .40,p >.10). foragingefficiency and reductions in diet breadth

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 419 duringcertain periods of the late Holocene. These withoutinterruption into the protohistoric period. trendsclosely mirror model-derived simulations Time-sensitiveprojectile points indicate sporadic of variationin climaticseasonality, paleoenvi- use duringearlier portions of theHolocene, but ronmentalproxy data on seasonality,and paleon- evidencefor human occupation from the terminal tologicalevidence for dramatic Holocene shifts in Pleistocenethrough the middle Holocene is ex- artiodactyldensities in theregion. Although fur- ceedinglyrare. Following convention for the area, theranalyses are plannedinvolving artiodactyl ourdiscussion of the LBB archaeologicalrecord skeletalpart representation and birdtaphonomy, dividesmaterials into three discrete time periods, extantdata stronglysupport the hypothesis that necessitatedby thefact that many assemblages climaticseasonality played a majorrole in con- can be datedonly with reference to projectile- trollingartiodactyl densities across the Holocene pointchronologies that are tiedto theseperiods in theBonneville Basin andthat human foraging (see Cannon2010). Theseinclude the Middle Ar- efficiencyand diet breadthresponded accord- chaic period,which in thecase of theLBB in- ingly.Moreover, that such a widevariety of mea- corporatesmaterials dating to betweenabout suresvary in a predictablefashion with the body 3,000 and 1,300B.R (comprisingonly approxi- size-basedArtiodactyl Index stronglysuggests matelythe latter half of the Middle Archaic period thatthe body size-return rate relationship is robust as thisperiod is recognizedthroughout the upper inthis case andthat the Artiodactyl Index is a use- HumboldtRiver region as a whole);the Maggie fulmeasure of foraging efficiency. Creekphase, spanning the period between about 1,300 and 650 B.R; and theEagle Rock phase, BoulderBasin Little whichranges from 650 B.R to thetime of Euro- Insofaras the climaticchanges evident in the Americancontact. The Eagle Rockphase is byfar BonnevilleBasin paleoenvironmental record also thebest representedin theLBB excavatedsite characterizedadjacent regions of the northern sample,while excavated Maggie Creek and Mid- GreatBasin, we anticipatesimilar trends in ar- dle Archaicassemblages are less abundant. tiodactyldensities and huntingefficiencies in The MaggieCreek phase is closelyaligned in thosesettings. One suchsetting is theLittle Boul- timewith one of thelate Holocenespikes in ar- derBasin (LBB), which,though occupied over a tiodactyldensities that are evidentat the Bon- shorterspan of timelimited primarily to thelate neville Basin sites of Hogup and Homestead Holocene,provides a dataset comparable to that caves. Indeed,this period of about1,300 to 650 fromHogup Cave bothin thevariety of archaeo- B.R was a timewhen climatic conditions through- logicalevidence that can be marshaledand in its outmuch of the Great Basin appearto havebeen consistencywith regional paleoenvironmental quite favorable for both human foragers and their patterns.The LBB is locatedapproximately 300 large mammalprey (see overviewin Grayson kmto the west of Hogup Cave inthe upper Hum- 2006). Wintertemperatures were likely elevated boldtRiver drainage (Figure 3). Researchcon- duringthis period,and summerprecipitation ductedin thecontext of culturalresource man- likelyreached its late Holocene maximum due to agementin thisarea has resultedin whatmay be morefrequent incursions of monsoonalstorms. thedensest concentration ofprofessionally exca- Bothof theseconditions should have resulted in vatedprehistoric archaeological sites ( n = -50) increases in artiodactylpopulation densities the LBB anywherein theGreat Basin (see Cannon2010). (Broughtonet al. 2008). By contrast, These sitesare predominatelyopen-air artifact MiddleArchaic and Eagle Rockassemblages date to scatters,many with numerous hearth features, to periods(3,000 to 1,300B.R and 650 B.R less-fa- whichappear to havebeen occupiedon a short- contact,respectively) characterized by We termbasis by mobile hunter-gatherers who passed vorableclimatic conditions for artiodactyls. indicesof throughthe area during the course of seasonal for- can thuspredict that archaeological be agingrounds. foragingefficiency in theLBB should higher The extensiveradiocarbon record from the duringthe Maggie Creekphase thaneither the or the subse- LBB suggeststhat sustained human occupation of precedingMiddle Archaic period thearea began around 3,000 B.R and continued quentEagle Rockphase.

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 420 AMERICANANTIQUITY [Vol.76, No. 3, 2011 Table6. LittleBoulder Basin Vertebrate Faunal Data by Time Period. Numberof Sitesor SiteLoci Total withFaunal ArtiodactylLagomorph Artiodactyl Vertebrate Period/Phase Material NISPa NISPb Index NTAXA NISPC EagleRock 14 326 640 .34 9 1013 MaggieCreek 3 423 15 .97 5 446 MiddleArchaic 2 2 29 106 7 83 includesspecimens identified toartiodactyl taxaas wellas specimensnottaxonomically identifiable butidentifiable tothe sizeclasses "large mammal" and "very large mammal". includesspecimens identified tolagomorph taxa as wellas specimensnottaxonomically identifiable butidentifiable tothe sizeclasses "small mammal" and "medium mammal". includesonly specimens identified totaxon; specimens identified only to size classes are excluded.

A summaryof faunaldata, including Artio- anddiet breadth changed in tandem in the manner dactylIndex values, for the three LB В timeperi- predictedby theprey model. Though we do not ods is presentedin Table 6. TheArtiodactyl Index evaluatethese differences inrichness statistically, is highestfor the Maggie Creekphase, and dif- we do notethat they do notappear to be driven ferencesin this index among the three periods are solelyby sample-size effects: fewer taxa are pre- 2 highlysignificant ( X = 503.6;df= 2,p < .001; all sentin the Maggie Creek phase assemblages than adjustedstandardized residuals fall beyond two in theMiddle Archaic assemblages, even though standarddeviations). This is consistentwith the theMaggie Creek sample is muchlarger than the predictionthat the favorable climatic conditions of MiddleArchaic one. theMaggie Creek phase led toincreased foraging A virtuallyidentical pattern occurs in thetax- efficiency.4And, as at HogupCave, several addi- onomierichness of plant resources. This is shown tionallines of evidenceare availablefrom the in Table 7, whichpresents data derivedfrom LBB tofurther test this prediction, supplementing counts of charredseeds fromradiocarbon-dated thetest provided by thebody size-basedArtio- hearths.Aggregate plant richness is lowestfor dactylIndex. theMaggie Creek phase, and this cannot be purely One such line of evidenceis the taxonomie an effectof samplesize sinceaggregate sample richnessof bothfaunal and macrobotanicalas- size is equal forthe Maggie Creek phase and semblages.As notedabove, a basicimplication of MiddleArchaic period; instead, this result is likely theprey model is thatdiet breadth should expand reflectinga narrowing of dietbreadth during the as ratesof encounterwith high-return resources MaggieCreek phase. At thelevel of theindivid- decline,and such an expansionshould be re- ual hearthfeature, as illustratedin Figure 10, flectedin an increasein taxonomie richness. Ver- plantrichness increases with increasing sample tebraterichness data from the LBB arepresented size forthe Middle Archaic period and the Eagle in Table 6. A comparisonof regressionsof Rock phase butdoes notdo so forthe Maggie NTAXA on samplesize is notuseful in thecase Creekphase, again suggestingthat diet breadth of theLBB data,as it was above in thecase of was narrowestduring this latter period. The dif- HogupCave, becausethe very small number of ferencesin regressionslope shownin Figure10 assemblagesfrom the earliest two timeperiods arenot statistically significant (F = .85; df= 2, p precludemeaningful regression analyses. Instead, = .434), butthey are consistentwith the pattern simpleaggregate NTAXA values,calculated by thatis tobe expectedgiven the Artiodactyl Index poolingthe assemblagesfrom each periodand andvertebrate richness data presented above. Col- countingthe total number of taxapresent in the lectively,these lines of evidencefrom the LBB aggregatesample from each period,are consid- faunaland floral data present a coherentpicture of ered.As expected,aggregate NTAXA is lowest highforaging efficiency and narrowdiet breadth duringthe Maggie Creek phase and higher during duringthe span between 1,300 and 650 B.R, with theMiddle Archaic period and theEagle Rock lowerforaging efficiency and broader diets both phase,suggesting that artiodactyl encounter rates beforeand after this period.

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 421 Table7. LittleBoulder Basin Macrobotanical e - • EagleRock Databy Time Period. EagleRock 5. 9 О MaggieCreek MaggieCreek Numberof Middle Archaic Radiocarbon-dated _ 4. « MiddleArchaic "wс Featureswith TotalNumber of °-SS Macrobotanical Macrobotanical < 3■ • О • • Period/PhaseMaterial NTAXA Specimens и- ..■••■•"l!-- Z 2- • - EagleRock 38 16 352 ф MaggieCreek 9 5 56 ~ 1- •"♦<>O o •" MiddleArchaic 14 8 56 o-I , , , , , , , , 0.000.25 0.50 0.75 1.00 1.25 1.50 1.75 2.00 2.25 Table8. LittleBoulder Basin Projectile Logn-Macrobotanical Specimens PointCounts by Time Period. Figure10. Relationships between macrobotanical richness and NumberofTemporally Numberof samplesize for each LBB time period. DiagnosticProjectile Projectile PointsReported from Pointsper 1.25 - Period/PhaseExcavated Sites CalendarYear 100 <^>Artiodactyl Index <$> Rock 207 .35 • ProjectilePoints/Year Eagle - MaggieCreek 618 1.03 0.75 - MiddleArchaic 305 Л6 I 0.50 Note: Counts ofElko series points, which were likely used ¿ 0.25- boththe Middle Archaic andthe Creek

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 422 AMERICANANTIQUITY [Vol.76, No. 3, 2011 Table9. LBBGround Stone Data by Time Period. Numberof Ground Numberof Shape Stone GSArtifacts Shape Shape Modification Period/PhaseContext Artifacts Area(m2) perm2 Modified Unmodified Index EagleRock Surface 4 7822 .00051 3 7 .30 Excavation 6 860 .00698 Total 10 MaggieCreek Surface 0 30741 .00000 0 4 .00 Excavation 4 340 .01176a Total 4 MiddleArchaic Surface 8 82681 .00010 5 39 .lib Excavation 40 291 .13746 Total 48 Note: Includes only ground stone artifacts such as manos,metates and pestles that likely served a food processing function. aAreais not reported forone excavation block, so this is a maximumestimate. bOfthe 48 Middle Archaic ground stone artifacts, 4 are not reported ina mannerthat allows determination ofwhether their shapehas been modified.

Two such measuresof investmentin grinding amountof timespent manufacturing food-pro- stonescan be calculatedfor the LBB (Table 9). cessingimplements. This index varies among the The firstis basedon theabundance of ground- threeLBB timeperiods in the manner that would stoneartifacts, which, consistent with the pat- be predictedbased on thetech investment model, ternsdescribed to thispoint, is lowestfor the thoughdue to smallsample sizes, the differences 2 Maggie Creek phase. Absolute artifactabun- arenot statistically significant ( X = 3.07,df= 2, dances,however, are somewhat problematic as an p = .216). Despitethe lack of statisticalsignifi- indicatorof technologicalinvestment since they cance,however, the shape modification index and areas mucha functionof the amount of effort that ground-stoneartifact density collectively present archaeologistshave expended at sitesof a given a consistentpicture of lower investment ingrind- age.To controlfor this factor, the ground-stone ar- ing technologyduring the Maggie Creekphase tifactsfrom each timeperiod can be subdivided thaneither before or afterthis time (Figure 12). intothose from surface and subsurfacecontexts, The ground-stoneshape modificationindex andartifact counts can thenbe normalizedeither (Table 9) also tracksaggregate plant richness by thetotal surface area of investigatedsites or (Table7) quitewell, suggesting that, in thiscase, siteloci, in thecase ofsurface artifacts, or by the thepatterns observed in the ground stone data are totalarea of excavationunits, in thecase of arti- reflectingpatterns in foragingefficiency and the factsfrom excavation.5 When thisis done,the breadthof the plant component of the diet. densityof ground-stoneartifacts from surface LittleBoulder contextsvaries in the manner predicted. The den- Basin: Summary sityof ground-stoneartifacts from subsurface Additionallines of evidencereported elsewhere contextsdoes notdo so, butbecause the area of (Cannon2010), particularlyevidence relating to oneMaggie Creek phase excavation block cannot hearthfeatures (see also Brightet al. 2002), pro- be determinedfrom the relevant excavation report, videfurther support for the conclusion that can be theexcavation density value derived for this phase derivedfrom the pattern in theArtiodactyl Index maysubstantially overestimate the true value. thatartiodactyl encounter rates and foraging effi- The secondmeasure of investment in ground- ciencyduring the late Holocene in the LBB were stonetechnology that can be derivedis a "shape highest,and diets narrowest, between about 1,300 modificationindex." This index reflects the abun- and650 B.P. It sufficesto sayhere, however, that danceof ground-stoneartifacts that are described a varietyof independentmeasures - including in excavationreports as beingformally shaped in vertebrateand plant taxonomie richness, projec- some mannerrelative to thosethat are described tilepoint frequencies, and both the density and the as notbeing so shaped,and it shouldreflect the degreeof modification ofground-stone artifacts -

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 423 1.25 uniformlyshow significant positive correlations О Groundstone/m2 ^ 1.00-ф ShapeModification Index| betweenprey size andpost-encounter return rate, 0.75- withjust a couple of exceptionsthat occur in 05°" cases whereonly a narrowrange of smaller-sized (Л8 25- preyare considered. Yet, as withall assumptions ¿Ф 0 Гоз О ОО - thatare incorporatedinto the application of any thatthe I -025 - # foragingmodel, we emphasize body - О size-returnrate rule of thumbshould be -0.50- always treatedas a hypothesisto be testedrather than as -0.75■ О an absoluteor invariantclaim about past human И.00 -1 1 1 « 1 MiddleArchaic MaggieCreek EagleRock foragingbehavior and prey choice. And because AnalyticalPeriod deviationsfrom the general body size-return rate do occur, devia- 12. ofLBB stoneartifacts from sur- relationshipcertainly potential Figure Density ground on a basis. facecontexts and ground stone shape modification index tionsmust be considered case-by-case (standardized)bytime period. In thisregard, we notethat there are many examples of archaeologicalapplications of foraging theoryin whichconsiderable attention has been are all consistentwith this conclusion. Thus, as devoted to such potential deviations (e.g., withthe Hogup Cave example,this case high- Broughton2004; Stineret al. 2000; Ugan2005a), of lightsthe usefulness of a bodysize-based abun- andwe also emphasizethat our understanding danceindex like the Artiodactyl Index as partof the variablesinfluencing them continues to be a comprehensiveeffort to track changes in forag- enhancedby detailedanalyses of contemporary ing efficiencyand importantrelated behavioral foragers(e.g., Bird et al. 2009). variables.The degreeof consistency among all of Forartiodactyls and lagomorphs, taxa of par- thevarious lines of evidenceconsidered in both ticularimportance in NorthAmerican archaeo- to exhibit examplesalso stronglysuggests that the body faunas,the available data showthem size-basedArtiodactyl Index is a usefulindicator dramaticallysignificant differences in return - thecosts of of changes in prehistoricforaging efficiency, rates evenin data sets incorporating here whichin thiscase appearsto be tightlylinked to failedpursuits. Further, modeling presented climaticvariability. suggeststhat it is highlyunlikely that pursuit success forlagomorphs could ever have been thanthat for to the extent Conclusion greater artiodactyls thatthis might have offset the difference in return in size Theuse of foraging theory to address issues related rateassociated with the difference body Previous toprehistoric foraging efficiency and diet breadth betweenthe two orders. archaeological haveused the size-return representsone of the more prolific areas of the ap- applicationsthat body forthese taxa are thus plicationof evolutionary ecology in archaeology. raterule of thumb partic- The successof this approach has beenenabled by ularlywell supported. mostof the de- thederivation of proxy measures of key model pa- Finally,we also emphasizethat a in appli- tailedanalyses of archaeofaunalvariability that rameters, requirement archaeological - cations.One of the most widely applied such prox- haveused the body size-return rate relationship indexor ies has beenthe use ofbody size as a measureof operationalizedthrough an abundance - have notrelied on thisrela- preyrank, and research drawing on thefoundation similarmeasures thesestudies have em- providedby thisproxy has had far-reachingim- tionshipalone. Instead, linesof rich- plicationsfor our understandingof manyother ployedmultiple evidence,including aspectsof human behavior and evolution. nessand diversity indices, body-part representation, of in- The use ofthe body-size proxy continues to be age structure,and measures technological Our of faunal, supportedby theavailable ethnographic and ex- vestment. analyses archaeological derivedreturn-rate data, informa- floral,and toolassemblages from the Bonneville perimentally of tionthat we have summarizedhere. These data and LittleBoulder basins represent examples

This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 424 AMERICANANTIQUITY [Vol.76, No. 3, 2011 thiskind of approach. In eachof these analyses we thePrescott Region, edited by Marvin D. Jeter,pp. have atleast half a dozendiet-breadth in- 339-367.Anthropological Research paper No. 1 1 .Arizona employed StateUniversity, Tempe, Arizona. dices,each of whichbehaves as predictedfrom a 1979 FactorsInfluencing theArchaic Pattern ofAnimal Uti- considerationofpaleoclimatic data from the north- lization.Kiva 44:219-235. 1982A Diachronic Prehistoric easternGreat Basin. In turn,the fact that such a di- Analysisof AnimalEx- ploitationatVentana Cave. Ph. D. Dissertation. Department versearray of archaeological variables vary in tan- ofAnthropology, ArizonaState University, Tempe, Arizona. demboth with each otherand withpaleoclimatic UniversityMicrofilms, AnnArbor. 1986Middle Archaic Animal Utilization inSouth Central variablesstrongly suggests that overall in patterns Arizona.InPrehistoric Hunter Gatherers ofSouth Central the archaeologicalrecord are reflectingclimati- Arizona:The Picacho Reservoir Archaic Project ,edited by callydriven variability in foragingefficiency and FrankE. Bayham,D. H. Morris, and M. S. Shackley,pp. dietbreadth. more for 315-340. Arizona State University Anthropological Field Perhaps important purposes Studies13. Tempe, Arizona. ofthis paper, the fact that the conclusions that can 2010Effects ofa SedentaryLifestyle onthe Utilization of be derivedfrom a body-size-basedabundance in- Animalsinthe Prehistoric Southwest. InEvolutionary Ecol- dex are so consistentwith conclusions derived ogyand Archaeology : Applications toProblems inHuman EvolutionandPrehistory, editedby Jack M. Broughton and fromso manyother lines of evidence attests to the MichaelD.Cannon, pp.1 59-1 71 . UniversityofUtah Press, robustnessof the body size-return rate relationship. SaltLake City, becausethe size ofa itemis Beaton,John, M. Moreover, body prey 1973 TheNature of ofMollusk oneof AboriginalExploitation Pop- thefew invariant referent properties of prey ulationsinSouthern California. Master's thesis, Department selectionin archaeologicalcontexts, this attribute ofAnthropology, University ofCalifornia, LosAngeles. and its to rankcan now even Beck,R. Kelly relationship prey 201 0 TheMolecular Genetics of Choice: Ancient more Prey Using confidentlybe usedand employed to maxi- DNAto Infer Prehistoric Population Histories. California mize theinferential potential of any prehistoric Archaeology1:253-268. dataset. We therefore conclude that, when used Betts,Matthew W.,and T. Max Friesen ju- 2006 Returnsfrom anInexhaustible Re- as one DecliningForaging diciously componentof broaderarchaeo- source?Abundance Indices and Beluga Whaling inthe West- logicalanalyses geared towards testing hypotheses ernCanadian Arctic. Journal ofAnthropological Archae- 25:59-81. derivedfrom foraging theory, the ology body-sizeproxy Bird,Douglas W., Rebecca Bird,and Brian F. for rankmust be considered Bliege Codding prey morereliable 2009In Pursuit ofMobile Prey: Martu Hunting Strategies thantenuous, and more robust rather than fragile. andArchaeofaunal Interpretation. American Antiquity 74:3-30. Acknowledgments.Wethank Andrew Ugan, Lisa Nagaoka, Bird, Douglas W., and James F.O'Connell DouglasBird, Brian andtwo reviewers 2006Behavioral Ecology and Archaeology. Journal ofAr- Codding anonymous Research14:143-188. forhelpful comments onthe manuscript. Wealso thank Bill chaeological Fawcettofthe Bright,Jason, Andrew Ugan, and Lori Hunsaker U.S.Bureau of Land Management, ElkoDis- 2002The Effect of Timeon Subsistence TuscaroraField Handling Tech- trict, Office,for his insights into and support nology.World Archaeology 34:1 64- 181. ofthe Little Boulder Basin research reported here. Broughton,Jack,M. 1994a Late Holocene Resource Intensification inthe Sacra- mentoValley, California: TheVertebrate Evidence. Jour- ReferencesCited nalof Archaeological Science 21:501-514. 1994b Declines inMammalian C.Melvin ForagingEfficiency During Aikens, theLate Holocene, San Francisco Bay. Journal ofAn- 1970 HogupCave. University ofUtah Anthropological Pa- thropologicalArchaeology 13:371 -40 1 . pers93. 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This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions Broughtonetal.] PREYBODY SIZE AND RANKING INZOOARCHAEOLOGY 425 nithologicalMonographs 56.Washington, D.C. Nevada.SWCA Cultural Resources Report No. 2009-87 Broughton,Jack,M., and Frank E. Bayham SWCAEnvironmental Consultants, SaltLake City. Sub- 2002Showing Off, Foraging Models, and the Ascendance mittedtoU.S. Bureau ofLand Management, ElkoDistrict, ofLarge Game Hunting inthe California Middle Archa- TuscaroraField Office, Elko, NV. BLM Project No. BLM ic.American Antiquity 68:783-789. 1-2505.Forthcoming inBLM Nevada Cultural Resource Broughton,JackM., David Byers, Reid Bryson, William Eck- Series(http://www.blm.gov/nv/st/en/prog/more-pro- erle,and David B. Madsen grams/cultural-resources/reports.html). 2008Did Climatic Seasonality Control Late Quaternary Ar- Cannon,Michael D., and Jack M. Broughton tiodactylDensities inWestern North America? 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This content downloaded from 155.97.85.112 on Wed, 26 Nov 2014 13:19:17 PM All use subject to JSTOR Terms and Conditions 428 AMERICANANTIQUITY [Vol.76, No. 3, 2011] dicesand Foraging Returns. Journal ofArchaeological Sci- carefulattention tolocally derived paleoclimatic data.And cli- ence28:1309-1321. maticconditions were, of course, not the only factors influ- Ugan,Andrew, Jason Bright, and Alan R. Rogers encingthe prehistoric abundances ofartiodactyls inwestern 2003When isTechnology Worth theTrouble? Journal ofAr- NorthAmerica. Despite generally favorable environmental chaeologicalScience 30: 1 3 1 5-1 329. TheodoreE. conditionsforartiodactyls across many stretches ofthe late White, incertain human 1952Observations onthe Butchering Technique ofSome Holocene, contexts, huntingpressure ap- AboriginalPeoples: I. American Antiquity 17:337-338. pears to have ultimately overtaken them, causing substantial 1953 AMethod ofCalculating theDietary Percentage ofVar- populationdeclines. Such anthropogenic depressions have iousFood Animals Utilized byAboriginal Peoples. Amer- nowbeen documented insome detail in several areas of west- icanAntiquity 18:396-398. ernNorth America. We also emphasize that variation inar- Wilson,Davis S. chaeologicalsitefunction orthe role that sites played inthe re- 1976Deducing the Energy Available inthe Environment: gional settlement-mobility systemcan, of course, influence the AnApplication ofOptimal Foraging Theory. Biotropica of choiceand andaffect variation in 8:96-103. energeticsprey transport, Bruce thetaxonomie composition ofarchaeological faunas, inde- Winterhaider, of trendsinthe 1977 Foraging Strategy Adaptations ofthe Boreal Forest Cree: pendentany temporal abundancesofartio- AnEvaluation ofTheory andModels from Evolutionary Ecol- dactylson past landscapes (Bayham 1982; Broughton 1999, ogy.Ph. D. Dissertation, Department ofAnthropology, Cor- 2002; Cannon 2003). A shift inthe regional settlement pattern nellUniversity. University Microfilms, AnnArbor. inwhich the function ofa site changed from a residential base 198 1 ForagingStrategies inthe Boreal Forest: An Analysis to a huntingcamp, would, for instance, have obvious implica- ofCree Hunting and Gathering. InHunter-Gatherer For- tions for variation inthe relative abundances oflarge game agingStrategies : Ethnographic andArchaeological Analy- skeletal elements orhunting tools recovered from its sedi- edited BruceWinterhaider andEric Alden ses, by Smith,ments (see Bayham 1982; Byers and Broughton 2004). And in- pp.66-98. University ofChicago Press, Chicago. Steve sofaras thereis a greaterlikelihood that large-packaged re- Wolverton, sourceswill be field 2005The Effects ofthe Hypsithermal onPrehistoric For- processed,weanticipate killsites to be agingEfficiency inMissouri. American Antiquity 70:91-106. biased towards the remains oflarge-sized prey, regardless of Wolverton,Steve,Lisa Nagaoka, Julie Densmore, andBen Fullerton when they were deposited. 2008White-tailed DeerHarvest Pressure andWithin-bone 3. Tomaintain consistency with the published literature NutrientExploitation During the Mid-to-Late Holocene in (e.g.,Broughton etal. 2008; Grayson 2006; Madsen 2000), un- SoutheastTexas. Before Farming 2008: 1-23. calibratedradiocarbon years before present isthe time-scale Yesner,David R., and Jean S. Aigner usedhere. 1976 ComparativeBiomass Estimates andPrehistoric Cul- 4.Consistent trends inthe Indexalso occur tural ofthe Southwest Umnak Aleutian Artiodactyl at Ecology Region, othersites with faunal inthe Islands.Arctic Anthropology 8:91-112. well-reportedassemblages upper Young,D. Craig HumboldtRiver region. At James Creek Shelter, located ap- 2004Geoarchaeology. InArchaeological Excavations atPie proximately25km to the southeast ofthe LBB, the abundance Creekand Tule Valley Shelters, Elko County, Nevada ,edit- ofartiodactyls relative tolagomorphs isalso highest indeposits edby Kelly R. McGuire, Michael G. Delacorte, andKim- datingtothe Maggie Creek phase (data in Grayson 1990). At berlyCarpenter, pp.39-53. Nevada State Museum An- PieCreek Shelter, located approximately 65km northeast ofthe thropologicalPapers No. 25. Carson City, Nevada. LBB,the Artiodactyl Index is highest indeposits that date to betweenabout 3,960 and 2,740 B.P. (Carpenter 2004; see dis- cussionofthe ofthis site in Cannon This Notes dating 2010). largely pre-datestheearliest faunal assemblages from the LBB but, no- tothe earlier ofthe two late Holocene 1.This formulation assumes for simplicity's sakethat av- tably,corresponds spikes timedoes not differ between successful andun- inartiodactyl abundance observed atHogup and Homestead eragehandling Caves.The successfulpursuits. ArtiodactylIndex then declines atPie Creek Shel- terin thatare coevalwith the LBB 2.Insofar asparticular spatiotemporal contexts the deposits roughly MiddleAr- during chaic ThePie Creek Shelter faunal data are un- earlyand middle Holocene were characterized bymore equable assemblages. climate,forinstance, alsohave den- fortunatelynotuseful for exploring trends inartiodactyl relative theymay supportedhigher abundance the sitiesof artiodactyls. Primary migration corridors also during MaggieCreek and Eagle Rock phases may becausematerials from these are havecontained relatively high densities (seasonally) ofgame phases stratigraphicallyco- duringthese times. In such contexts, increases inthe mingledatthe site (see Young 2004:45^-6). propor- 5.Due to lowrates of the tionalrepresentation ofartiodactyls would not be anticipated very depositionduring Holocene, atthe middle-late Holocene transition. buriedarchaeological materials inthe LBB generally occur within Aswe discuss elsewhere etal. 2008: 193 1 sev- thefirst 20cm below surface, andexcavations routinely proceed (Broughton ), no thanthis. eralrecords seem to show just this pattern. Westress too that deeper Therefore,excavation unitarea, rather than vol- ume,isan measureofexcavation effort. artiodactylabundances appear to have been highly variable appropriate throughtime within thelate Holocene, asthe Bonneville Basin paleontologicalandarchaeological records soclearly suggest. Thus,generating predictions about variation inartiodactyl Submitted January 29,2010; Revised March 18, 2010; abundancesatany particular spot on the landscape requires Accepted March 19, 2010.

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