using evolutionary archaeology and evolutionary ecology to explain cultural elaboration: the case of...

8/7/2019 Using Evolutionary Archaeology and Evolutionary Ecology to Explain Cultural Elaboration: The Case of Middle Ohio

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NORTH AMERICAN ARCHAEOLOGIST, Vol. 31(2) 119-154, 2010

USING EVOLUTIONARY ARCHAEOLOGY ANDEVOLUTIONARY ECOLOGY TO EXPLAIN CULTURALELABORATION: THE CASE OF MIDDLE OHIO VALLEYWOODLAND PERIOD CEREMONIAL SUBSISTENCE*

KEVIN C. NOLAN

STEVEN P. HOWARD

Ohio State University, Columbus

ABSTRACT

We propose a Darwinian evolutionary model for the development and dis-appearance of Woodland period cultural elaboration in the Ohio River Valley, and specifically the climax of this behavior known as the Hopewellphenomenon. We combine aspects of evolutionary archaeology and evolu-tionary ecology to provide a model that (1) has testable empirical conse-quences, and (2) specifically addresses the historical context of developmentof the phenomenon being explained. Our model builds on Smiths (1987)coevolutionary scenario for the development of symbiosis between EasternAgricultural Complex (EAC) crops and the human populations exploitingthem. After presenting our model, we explore the empirical consequencesof ceremonial subsistence and the types of data that would be required to testthe model. In our discussion of the extant data we also contrast our scenariowith a competing evolutionary explanation for the Hopewell phenomenon:the waste hypothesis. The limited available evidence is in line with theempirical expectations of the ceremonial subsistence model and is, to varying

*An earlier version of this article (Nolan and Howard, 2007) was presented at the 53rd annualmeeting of the Midwest Archaeological Conference at Notre Dame, Indiana.

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degrees, at odds with the empirical expectations for the waste explanation.Our ceremonial subsistence model offers a testable alternative that is notfalsified by the extant data. However, we recognize the problem is far fromsolved. Our primary aim is to promote renewed theoretical discussion of theissues raised herein and to encourage new problem-oriented research toprovide empirical evidence to test both explanations.

INTRODUCTION

Evolutionary archaeology and evolutionary ecology approaches to archaeologyare increasingly being recognized as complementary (e.g., Gremillion, 2006;Lipo and Madsen, 1998; Lyman and OBrien, 1998; see also Laland and Brown,2002). We contribute to this growing trend by proposing a Darwinian evolutionarymodel that mixes elements of the two approaches to explain a particular case of cultural elaboration. We also provide a fresh assessment and partial critique of analternative explanation for the same cultural phenomenon: the waste hypothesis(Dunnell, 1996c; Dunnell and Greenlee, 1999; Madsen et al., 1999). By con-trasting the empirical consequences of both explanations and examining theavailable evidence, we identify an apparent lack-of-fit for the waste hypothesisand describe the types of data required to fully evaluate both hypotheses. Dueto the current coarseness of both models and the limited available data, we areunable to fully reject either explanation.

For our case study, we focus on the development and disappearance of Wood-land period cultural elaboration, specifically on the Ohio Hopewell (ca. 50 BC

AD 400) zenith of this elaborative behavior. Both evolutionary archaeology andevolutionary ecology have been used to explain monumental architecture andcultural elaboration such as that exhibited by the Ohio Hopewell. Each approachhas been successful at presenting plausible hypotheses for elaborative behavior in various circumstances, and evolutionary archaeology has been appliedspecifically to the Ohio Hopewell phenomenon (Dunnell, 1996d; Dunnell andGreenlee, 1999). We feel that neither of these general explanations are a completeexplanation of Ohio Hopewell cultural elaboration. With a combination of aspects of evolutionary archaeology and evolutionary ecology, we propose ahypothesis that is plausible and empirically testable. We hope this article willspur theoretical discussion and empirical evaluation of explanations such as theones discussed herein.

PREVIOUS EVOLUTIONARY MODELS OFELABORATIVE BEHAVIOR

Both evolutionary archaeologists and evolutionary ecologists note that culturalelaboration is seemingly wasteful from a fitness standpoint, and both attempt todemonstrate that it can be accounted for via individual-level fitness maximizing

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strategies. The waste hypothesis from evolutionary archaeology (e.g., Aranyosi,1999; Dunnell and Greenlee, 1999; Hamilton, 1999; Kornbacher, 1999) andcostly signaling from evolutionary ecology (e.g., McGuire et al., 2007; Neiman,1997; Smith et al., 2003) have been employed in a number of instances to provideevolutionary explanations for seemingly inefficient behavior. Both hypothesesdraw upon Darwinian Theory and evolutionary explanations developed inevolutionary biology for behavior of non-human species.

The costly signaling hypothesis suggests that an individual, by advertising their quality, increases their fitness, and that the more costly the signal, the more honest(because it is less likely to be faked). Contests between males, for example, cansignal to females which one is more able to provide for offspring. The concept hasbeen employed convincingly to explain individual behaviors, such as materialdisplay and risky hunting behavior (e.g., Bird and OConnell, 2006). Although ithasnt been applied to the Hopewell situation, costly signaling theory has beenused to explain monument building in other regions. Neiman (1997) suggeststhat Mayan elites were signaling their prowess to each other and prospectivefollowers through their erection of monumental architecture. In the case of theHopewell earthwork builders in the Middle Ohio Valley, there is no evidencethat any individual was ranked highly enough to be solely responsible for theconstruction of the earthworks. Therefore, it would be unclear exactly who wassignaling. To date, no one has proposed a version of costly signaling as anevolutionary explanation of the Hopewell phenomenon against which to evaluatethe evidence, so we focus our discussion and comparisons on the wastehypothesis, which has been applied to the Hopewell situation.

The waste hypothesis was originally formulated by Dunnell in 1989 todemonstrate the utility of an evolutionary archaeology approach (Dunnell, 1996d;see also Dunnell, 1999), using the elaborative behavior of the Ohio Hopewellas an illustrative example. Dunnell and Greenlee (1999) refined the argumentthat mound building behavior diverts energy from reproductive behavior andtherefore is wasteful, apparently reducing the fitness of the individualsinvolved. Dunnell (1999:246) argues that waste can be favored by naturalselection acting in the usual fashion in somewhat unusual circumstances. Within(unusually) unpredictable environments, seemingly wasteful behavior canactually benefit individual fitness by diverting energy into activities other thanproduction (and reproduction), thus reducing the numbers of offspring. Thiskeeps the population within the local carrying capacity during lean times (seeDunnell, 1996d:Figure 4.2).

It is widely known in the biological sciences that organisms will divert energyaway from reproductive effort in temporally variable environments to increaseeffective fitness (Dunnell, 1999:245; Madsen et al., 1999; see also Boyce andPerrins, 1987; Seger and Brockman, 1987). In such environments it pays toproduce offspring below the organisms maximum potential in order to raiseoffspring of bettercompetitive quality to the age of reproductive viability (Madsen

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et al., 1999:Figure 3). The logic is that organisms will sacrifice some expectedfitness so as to reduce their uncertainty, or variance, of fitness (Seger andBrockman, 1987:185). According to Seger and Brockman (1987) this is accom-plished by maximizing the geometric-mean fitness (cf. arithmetic mean; seealso discussion in Madsen et al., 1999). This same logic may be applied tohuman reproductive behavior. Madsen et al. (1999) used computer simulations todemonstrate that, under the Darwinian theory of natural selection, seeminglyinefficient reproductive regimes will persist and in fact increase in frequencyunder some circumstances. The simulations of Madsen et al. are re-illustrationsof the geometric-mean principle of Seger and Brockman (1987). However, inthe non-human cases that provide the theoretical basis for waste/bet-hedgingbehavior, energy diverted from reproduction is not invested in elaborativebehavior. Excess energy can be reinvested in parenting effort, which makesgood Darwinian sense.

Alternatively, below optimum reproduction in fluctuating environments issimply the best long-term bet to maximize individual lifetime reproductivesuccess (Boyce and Perrins, 1987). Variable environments select for reducedproduction of offspring only, not for cultural elaboration. The proponents of thewaste hypothesis acknowledge that there is no necessary causal connection(Madsen et al., 1999); however, they do not address how the linkage woulddevelop. For example, Dunnell and Greenlee (1999) focus only on how elabor-ative behavior may have functioned and came to an end, with no attempt topostulate how the linkage became established. If the waste hypothesis is tobe used to explain the origin (in addition to function, persistence, and cessation)of elaborative behavior it seems necessary to address how the link betweenfitness and elaboration becomes established. In other words, what is the historicalcontext within which the behavior(s) evolved?

Additional issues are raised by Madsen et al.s (1999) reframing of Dunnellsoriginal model. Dunnells original presentation of waste was in terms of a trade-off in consumption of a finite energy budget (Dunnell, 1996d:95-97).Madsen et al. recast the model in terms of bet-hedging ( sensu Seger andBrockman, 1987) changing the focus of the explanation. Seger and Brockman(1987) and Boyce and Perrins (1987)both heavily cited by Madsen et al.(1999)do not make any mention of a trade-off in energy allocation of thequalityvs.quantity variety in their discussions of reproduction below optimumlevels under conditions of generation-scale temporal environmental variation.In fact, Boyce and Perrins (1987) find little support for the cost hypothesis,

which involves the kind of trade-off invoked by Dunnell (1996d), in their study of Great Tit clutch size. The conditions that would select for the twotypes of scenarios are also different. The trade-off (cost) hypothesis posits thatbelow maximum reproductive effort is selected for when risk of adult mortalityincreases with clutch size (Boyce and Perrins, 1987:142). In contrast, the bet-hedging type of reproductive restraint is predicted when there is increased



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variability in survivorship of offspring (Boyce and Perrins, 1987:142; Seger andBrockman, 1987). Therefore, bet-hedging and Dunnells original presentation of waste are two theoreticallypossible evolutionary explanations for the regulationof reproduction under different sets of circumstances.

Madsen et al. (1999) attempt to merge Seger and Brockmans (1987) definitions2.4 and 2.5 of bet-hedging with Dunnells original trade-off concept, but this isproblematic. Dunnells use of the energy trade-off establishes a relationshipbetween waste (non-reproductive use of energy) and long-term reproductivesuccess in varying environments. Madsen et al. (1999) attempt to rely mainly onbet-hedging as defined by Seger and Brockman (1987) and observed by Boyceand Perrins (1987) to explain increased elaborative behavior via a reduction infecundity, without establishing the connection between the two. The theory reliedon by Madsen et al. (1999) is capable of accounting for decreased fecunditywithout cultural elaboration. Madsen et al. recognize this gap when they statethat the linkage between elaboration and reproductive effort is an evolved [read:historically contingent], not mechanical, association (Madsen et al., 1999:267).However, they do not speculate how the two become linked or why the additionof elaboration would be necessary when the biological bet-hedging theoryis sufficient to explain reduced fecundity in temporally variable environmentswithout this diversion of energy (a circumstance that would not leave much of an archaeological signature). Madsen et al. (1999) claim their simulations demon-strate that wasting phenotypes would increase in frequency. However, they aresimply re-illustrating the geometric-mean principle and make the unnecessaryassumption that cultural elaboration is linked with lower fecundity and further that it is causally related to reduced fecundity.

While we cannot rule out the possibility of regulating numbers through elabor-ation as Madsen et al. (1999) present it, our analysis demonstrates that more work is necessary to fully develop the bet-hedging model for cultural elaboration.The theoretical principles may still apply and the fit of the model to each case isempirical. More attention needs to be focused on the historically contingent set of circumstances under which elaboration would become linked with a bet-hedgingtype of regulation of production of offspring. Whether Dunnells version of waste or Madsen et al.s version bet-hedging/waste can explain a given set of archaeologicalcircumstances is an empirical rather than theoretical determination.

To clarify, the fact that Madsen et al. (1999) did not identify the empiricalhistorical circumstance that did or may have resulted in linking elaboration withfecundity does not invalidate the theoretical argument in the case that such a link

can be established. Indeed, their paper was wholly theoretical and therefore couldnot have identified a general historical case for the association between the two.However, this absence does make the particular application difficult. For thisreason we attempt to trace the historical trajectory leading to elaboration.

The line of reasoning presented above does not weaken thegeneral applicabilityof either the trade-off or bet-hedging models to explain patterns in prehistory (not



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limited to cultural elaboration). The overall logic of these explanations and their connection to theory is sound. Dunnells (1996d) original intent of highlightingthe applicability of evolutionary theory to the archaeological record is importantto this discussion; however, his 1989 paper did not critically consider evidence insupport of his example. It simply illustrates how the logic of well establishedevolutionary models could be applied to problematic cases in the archaeologicalrecord; a lesson we see as crucial.

Bet-hedging (with or without elaboration) may very well explain many thingsin the archaeological record. However, Madsen et al.s (1999) specific applicationto cultural elaboration does not seem to be sufficient. It is easy to imagine thata few families not participating in elaborative behavior, but still observing belowoptimal reproductive rates would be at least as fit as their wasting counterparts,if not more so.

We are not arguing that these versions of the waste hypothesis are notplausible evolutionary explanations in some scenarios. We merely hope to illus-trate by our theoretical analysis above and our presentation below that, giventhe currently available data, the case for either or both of these hypotheses asexplanation for the Ohio Hopewell phenomenon is less than complete. First, thelink between elaboration and increased survivorship has not been establishedby specific presentations of the waste hypothesis. Second, as we will show below,the empirical record evidences several departures for the expectations derivedfrom the waste hypothesis.

If the development of earthwork construction in the Middle Ohio Valleyis to be accounted for by the action of natural selection (though there isno necessary reason that this must be the case), then we argue that ascenario that provides a direct contribution to fitness is more likely. It iswith the intent of exploring this possibility that we present the ceremonial sub-sistence model.

CEREMONIAL SUBSISTENCE

In this section we layout our alternative hypothesis for the origin and disappear-ance of the Middle Woodland period (circa 50 BC AD 400) cultural elaboration.We focus the ceremonial subsistence model on the Ohio Hopewell groups of the Middle Ohio River Valley (though occasionally drawing on discussion of neighboring regions of necessity). Behavioral modifications leading to thedevelopment of the Hopewell climax (Middle Woodland) have their roots in the

Middle Archaic period (ca. 60003000 BC).In our presentation we rely on the culture-historical terminology predominantin the region (see Table 1). However, we should note that these terms are used invarious ways by various authors and in all cases they are largely common senseterms and in need of theoretical justification for their definitions (e.g., Hartand Brumbach, 2003). Most often the culture-historical terms are used to denote



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units of space-time. This is the conceptually most prohibitive use and we avoidthat here. For ease of understanding, we use the terms Archaic, Woodland, andtheir subdivisions as referents to blocks of time, not normative material packagesor cultural traditions (though the latter is their original meaning). We also,sparingly and regrettably, use the terms Adena and Hopewell to refer to aggregatepackages of interpreted behavior. This usage, we fully acknowledge, and hopebecomes clear to the reader, is at odds with our own perspective on the nature of the archaeological record and cultural change. We would note that even Dunnell(Dunnell, 1996d; Dunnell and Greenlee, 1999)a marked critic of the nature of extant, common-sense-based systematics in American Archaeologyemployedthese terms is his argument for the waste hypothesis. These terms have widepurchase in the region and are therefore retained here for efficiency of communication at the expense of precision. We hope these choices do not cloudout the nature of our argument.

As we proceed we will identify the types of evidence needed to evaluateboth the waste and ceremonial subsistence models and assess how each fitthe available data. Our goals in this article are to offer an alternative modelto compete with the waste model, and evaluate the veracity of both hypothesesagainst the available evidence. It is very important to note that currently there

is not enough evidence to fully evaluate either explanation, and we are not ableat this point to prove our hypothesis. However, in a scientific endeavor proof is not the goal ; we can either reject or fail to reject a given hypothesis. Thecurrent data and interpretations discussed raise questions about the applic-ability of the waste hypothesis and do not allow us to reject the ceremonialsubsistence hypothesis.


Table 1. Generic Time Periods for Eastern North American Prehistory a

Period Subperiod Dates

Archaic

Woodland

Late Prehistoric

EarlyMiddleLate

EarlyMiddleLate

AD 10001650

80006000 BC60003000 BC30001000 BC

1000 BCAD 1000100050 BC50 BCAD 400

AD 4001000

(Adena)(Hopewell)

aGenerally associated cultural packages are shown in parentheses after the dates. Notethat there is no necessary association between the cultural package and the dates of theperiod and that Adena and Hopewell are found to start at different times in different places.


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Historical Background for the Model

The basis for the ceremonial subsistence model is the development of theEastern Agricultural Complex (EAC), increased reliance upon food productionfrom the Middle Archaic period through the Middle Woodland period, and relatedchanges in settlement patterns (Smith, 1987, 1989, 1995; see also Braun, 1987).The ceremonial subsistence model begins with Bruce Smiths coevolutionaryscenario for the establishment of an association between human disturbed habitatsand abundance of edible plants.

Bruce Smiths Floodplain Weed Theory specifies how plant domestication andcultivation grew out of behavioral responses [e.g., increasing residential stabilityand modification of the environment] to particular features of the environment

(Gremillion 2002:492). This provides a historical context for the development of the symbiosis between anthropogenically disturbed habitats (domestilocalities)and increased local abundance of protodomesticates ( sensu Smith, 1987)and eventual domesticates. The Ohio Hopewell core falls squarely withinGremillions (2002:Figure 22.3) zone of developed pre-maize agriculture (seealso Smith,1989:Figure 1). We link the maintenance of disturbed habitats with thepractice of earthwork construction and posit that the continued evolution of EACplants takes place in association with, and perhaps partially within, the largegeometric earthworks for which Ohio Hopewell groups are so well known.

The Ohio Hopewell and the Rise and Fall of Elaboration

Most research into the monumental architecture and cultural elaboration of Ohio Hopewell has focused on proximal cultural causes. This has been a fruitfulavenue of research, recently producing numerous hypothetical, emic explanationsfor various aspects of Hopewell cultural elaboration. Byers (2004) postulates thatthe earthworks and their associated mortuary offerings are the result of worldrenewal cults trying to maintain balance within the cosmos. Others have focusedon the geometric and astronomical aspects of the earthworks, positing that manyof the sites served as calendars (e.g., Romain, 2000). Carr (2006) suggests that,in some cases, several earthworks in a region served differing, yet complementaryceremonial functions, each important to the regional populations in their ownright. Lepper (2006) even hypothesizes that long distance pilgrimages figuredinto the Hopewell ideology, making the earthworks important for people over large geographic ranges. While we acknowledge the significant contribution of these approaches to understanding Ohio Hopewell ideology and its role in cultural

elaboration, we do not address these hypotheses here because any of the variousemic (proximate) explanations may be valid, and none would necessarily beinconsistent with an overall evolutionary explanation (either bet-hedging/wasteor ceremonial subsistence). We are not concerned with emic aspects of the ritualsassociated with earthwork construction. We are concerned with explanation at aseparate and not exclusive level.



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The evolutionary approach asks different kinds of questions. Did the popu-lations that participated in cultural elaboration, for whatever emic reason, receivea fitness-related benefit in return for their costly investment of time and labor?How was the practice able to spread for a time, while seemingly less costlyalternatives were available? Could environmental, fitness-related changes haveled to the ultimate and abrupt abandonment of the practice across its entiredistribution? For our purposes, of more importance than the details of theceremony are the details of settlement and subsistence strategies.

There is much debate over the nature of the Early and Middle Woodlandperiod subsistence and settlement patterns (Cowan, 2006; Dancey and Pacheco,1997a, 1997b; Pacheco and Dancey, 2006; Weller, 2005; Wymer, 1996, 1997,2009; Yerkes, 2002, 2009). It is during the Early Woodland period that potteryfirst enters the record around 800 BC . Ceremonialism and trade became importantduring the Early Woodland with the most visible signs of this found in themore frequent construction of mounds and earthworks. Exotic materials becomeregular and more prominent members of the artifact inventory. The newceremonial and exchange activities increased in frequency and quantity duringthe Middle Woodland period with tons of soil and sediment moved to modifythe landscape and create large geometric enclosures, some containing multiplemounds (e.g., Bernardini, 2004; Greber, 1997).

The most well known aspects of the Early and Middle Woodland periods arethe nature of the large earthworks that dotted much of the Eastern Woodlandsand especially southern Ohio; however, there are many less well known smaller and less obtrusive earthen constructions not associated with the larger sites(e.g., Nolan, 2009, 2010; Nolan et al., 2008).

Even less is known about the habitation sites associated with the Middleand Early Woodland periods (e.g., Weller, 2005:7-10). Investigation of habita-tion locations is a relatively recent emphasis in Ohio Middle Woodland studies(Dancey and Pacheco, 1997a; Pacheco, 1996).

Settlement patterns largely were ignored for much of the history of archaeo-logical interest in Hopewell societies as researchers concentrated on investi-gating the major earthwork complexes. A renewed interest in settlement patternswas sparked in the late 1980s and 1990s by Dancey and Pacheco (Dancey, 1991,1992, 1998; Dancey and Pacheco, 1997a, 1997b; Pacheco, 1996, 1997). There areessentially two views of Middle Woodland period settlement systems and thereis ongoing and vigorous debate between the two camps. Some researcherscontend that the Hopewell groups that constructed the massive earthworks were

tribal societies. . . [that occupied] different sites . . . throughout theyear (Yerkes,2009:119; see also Cowan, 2006; Yerkes, 2002, 2003). On the other end of the debate, some researchers postulated that the Ohio Hopewell settlementpattern conforms to the Vacant Center/Dispersed Hamlet model first put forthby Olaf Prufer in 1965. The model holds that Hopewell family groups lived insmall, semi-permanent, homesteads dispersed around, but not within the vacant



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ceremonial centers (Dancey and Pacheco, 1997b). Vacant, as it is used byDancey and Pacheco, and Prufer, does not mean devoid of human activity, or archaeologically devoid of habitation debris. It simply means that there wereno permanent settlements within the ceremonial centers (Pacheco, 1996, 1997;Prufer, 1996). This point seems to be continually lost on critics of the model, someof whom consistently argue that the presence of any domestic debris at thesecenters contradicts the model (Griffin, 1996, 1997; Lepper and Yerkes, 1997;Yerkes, 2003:23).

What exactly qualifies as in the general vicinity of ceremonial centers(Pacheco, 1997:44), but not adjacent to such centers (Dancey and Pacheco,1997b:6) is not quite clear, and therefore the dispersed quality of the settlementpattern is difficult to assess with respect to what is expected for the model(Church and Ericksen, 1997:343). Pacheco et al. (2006, 2009) have recentlyexposed two such hamlets within the catchment of the Liberty Earthworks near Chillicothe, Ohio, clarifying to some extent the spatial relationship betweenadjacent hamlets, and between hamlets and earthworks.

The elaborate behaviors associated with Hopewell dwindle and disappear atthe end of the Middle Woodland period around AD 400 (Dancey, 1996; Dunnelland Greenlee, 1999). While the Late Woodland period in Ohio is little known(Seeman and Dancey, 2000), the early Late Woodland ( AD 400/500700/800)is generally characterized by large, sometimes fortified nucleated settlements(Church, 1987; Church and Nass, 2002; Dancey, 1992, 1998; Seeman andDancey, 2000). The late Late Woodland ( AD 700/8001000) is generally charac-terized by a return to small, dispersed settlements, though there are exceptions(Church, 1987; Church and Nass, 2002; Seeman and Dancey, 2000). Maizebecomes a major part of the subsistence economy circa AD 800 (Greenlee,2002; Hart, 1999:Figure 3).

The Model

The ceremonial subsistence model begins with the increasing residentialstability of Smiths (1987, 1995) hypothesis for the origin of the EAC. Followingthat trajectory forward in time, we expect relatively stable, long-term settlementsduring the period of development of the ceremonial subsistence behavior (Earlyand Middle Woodland). Using Bruce Smiths scenario as a point of departure for our model we find our expectations regarding the issue of Hopewell residential

stability in line with the Dancey-Pacheco model (Dancey, 1996; Dancey andPacheco, 1997a, 1997b; Pacheco, 1996a, 1996b; see also Prufer, 1996). Whileweighing in on this debate would be an unnecessary distraction from our task, some degree of permanence and regularity of land use is required by our model, but it does not depend on sedentary groups unflaggingly wedded toparticular habitation locations. We would point out that from an evolutionary



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perspective, spatial and temporal variability in group residence patterns wouldnot be unexpected.

Inevitably, resources surrounding long-term settlements become depleted andthe habitation must be relocated. As settlements are relocated in previouslyundisturbed habitats, the protodomesticates would spread with the people.However, the old habitation sites would not return immediately to mature forest.Old habitation sites would gradually move through the local successional stagesleading to mature communities. As Gremillion et al. (2008:395) point out [i]nthe humid temperate forests of eastern North America, the removal of maturetrees initiates a successional sequence in which colonizing annuals invade. . . .These weedy annuals persist only until the following year, when more shade-tolerant perennials and biennials establish themselves (Bazzaz, 1996:38-60;Runkle, 1985). This would provide additional patches of densely packed seedsand greens (e.g., Chenopoduim berlandieri [goosefoot], Helianthus annuus[sunflower], Iva annua [marshelder/sumpweed]), temporarily increasing thecarrying capacity of the local environment.

If humans continued to disturb the soil in the vicinity of these older habitationsites, the vegetation would remain in an artificially retarded successional stageprolonging the elevation of the local carrying capacity. Any activity, regardlessof emic intent, that functioned to delay the onset of mature plant communitieswould be favored in this way by natural selection. That is, if there is some reasonfor people to continue to visit the disturbed areas associated with their earlier habitations or cemeteries, then there would be an associated increase in thequantity of these annual seeds and therefore local carrying capacity.

A possible proximate cause of the human population returning regularly isto visit and/or inter their ancestors. Additionally, maintaining stands of nuts(Gardner, 1997) and/or productive, ecotone hunting grounds could serve asimpetus for maintenance of forest openings (both of these would also directlybenefit fitness). Even small levels of human disturbance in the area would helpmaintain the retarded successional state, resulting in maintenance of productivepatches of edible annual seeds and greens. Irrespective of the emic intent for continued human presence and activity at these previously disturbed sites, anyactivity that maintains an arrested state of succession will have the effect of increasing the abundance of edible resources on the landscape, and will thereforeincrease individual fitness and be favored by natural selection. Importantly,the activities that are maintaining disturbed areas need not be intended by theactors to increase productivity of the environment. This means that whether or

not the activity was an efficient way to accomplish the end is irrelevant. What isrelevant is whether or not performing that activity provides more resources thannot performing the activity, and whether or not performing that activity providesmore returns than contemporaneous alternatives.

As the colonizing EAC crops are low ranked resources, a high level of investment in managing their life cycle is not predicted except under times of



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food stress (Barlow, 2002; Winterhalder and Goland, 1997). As the density of the seed bearing annuals increases they would shift into what Winterhalder and Goland define as a low ranked, high density resource (1997:Figure 7.2).Winterhalder and Goland predict this would lead to local human populationdensity increases and a high risk subsistence system. Gradually, populationgrowth would apply stress to the subsistence system and increasing levelsof interference in the plants life cycles is expected (e.g., sowing, tending, culti-vating the soil).

Continued use of old cemeteries for the interment of the newly deceased and/or other ritual activities and maintenance of open landscapes (possibly associatedwith early mound and earthwork construction prior to or at the advent of theWoodland period) would thus have resulted in maintenance of populations of weedy plants with edible seeds with low energy investment. It must be remem-bered that the EAC crops are colonizing plants and do not require much (if any) tending to compete and mature in disturbed domestilocalities (Smith,1987, 1995). In this way activities associated with maintaining disturbed areaswould have had the effect of increasing the productivity of the local environment.With this mutual dependence established, any individual or family that investsenergy (for whatever proximate, emic intention) into maintaining areas of disturbance on the landscape will increase their resource base. This would affectnot only the abundance of crops, but also, indirectly, the availability of gameanimals in the vicinity. Deer are known to prefer open forest/forest edge situationsand are drawn to ecotones and open areas for their preferred foraging (e.g.,Barber, 1974:Table 9; Breitburg, 1992; Styles, 1981:Table 2). As the deer areunlikely to frequent current habitation sites, the maintenance of open areas awayfrom living humans will have the effect of reducing search times for this highlyvalued prey item and potentially buffering unsuccessful hunts with the bonus of abundant plant harvests at thesame location (see also Gremillion et al., 2008:397).

Therefore a selective advantage (e.g., increased fecundity, increased off-spring survivorship) is realized by any individual or family group that engagesin this type of maintained disturbance behavior. Relative to those who do notrevisit old disturbed areas this practice will result in more efficient returns onenergy expenditure, and, according to Winterhalder and Golands (1997) model,increased local population.

The particular fashion by which the disturbed area is maintained will be ahistorically contingent cultural behavior and may be inefficient as far as systemsof subsistence intensification are concerned; however, it does increase the density

of protodomesticates and the carrying capacity of the local catchment. The factthat the EAC seeds are readily storable makes it likely that the advantage tothese disturbers is to be realized most in the lean season. In fact, processing of small seeds during the lean season, when production effort had few competitors(i.e., low opportunity cost), is one of the likely reasons for their inclusion in thediet in the first place (Gremillion, 2004).



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There are very obvious implications for the archaeobotanical assemblagesassociated with this shift in subsistence. In fact, some of the expected changesalready have been documented in available data sets. For example, we expect anincrease through time in the ubiquity of EAC seeds and pollen at and near earthwork and habitation sites (see McLauchlan, 2003; Wymer, 1996, 1997; anddiscussion below). Braun (1987) identifies just such a pattern for the IllinoisValley during the Early and Middle Woodland period associated with trends inceramic vessel wall thickness.

Woodland Period Monumental Architecture

With the well established symbiosis and the selective advantage conferredon agents of disturbance, any set of behaviors that perpetuates this disturbance isbeneficial. During the Late Archaic, we expect increasing investment in main-taining open plots of land. As the EAC seeds continue to compete in and adapt tothe incipient agroecology ( sensu Rindos, 1980, 1984) created by these ceremonialsites and other open areas, selection will continue to favor those individuals(humans) that perpetuate these open areas. By the Early Woodland period, weknow that groups of people began to erect monuments associated with mortuarybehavior. Whatever the emic reason, the disturbances associated with this activitywould function to maintain an early stage of succession (e.g., see pollen diagramsin McLauchlan, 2003). The mortuary and ceremonial practices of the precedingperiod were sufficient to maintain modest increases in domesticate productivity,but the addition of moderate to large scale movement of earth increases the levelof disturbance and potentially the spatial scale of the disturbance maintained.This new variant ( sensu Boyd and Richerson, 1985) is not expected to appear everywhere at once; rather, it developed by chance in a few communities (perhapseven one). Eventually the practitioners of this variety of disturbance were moresuccessful at maintaining higher yields over long periods of time. Thus thepractice was favored by natural selection acting in the usual fashion (Dunnell,1999:246) in not so unusual a circumstance.

At this point the reader might ask: Why wouldnt the prehistoric incipienthorticulturalists simply plant gardens? Why build earthworks and maintainceremonial sites to indirectly increase subsistence yields? We expect that house-holds did maintain independent gardens. If people simply intended to maximizetheir long-term subsistence yield in the most direct fashion they would likelyskip the community-level ceremonial aspects of subsistence production (indeed,

we expect this eventually happened). However, individual intentions have littlerole to play in long-term evolutionary narratives. Emic intentions can serve ina proximate causal role for part of the historical sequence. The incipient farmersof the Ohio Valley did not intend to become agriculturalists (see especiallyRindos [1980:769-770] reply to commentators; also contributions to Price andGebauer, 1995).



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Evolution is historical and opportunistic. Natural selection acts on extantvariability; it does not forge it de novo . In our scenario, the ritual behaviorsassociated with maintenance of cleared space and eventually massive, regular movements of dirt would increase the return rate of foraging efforts therebydecreasing the level of time and energy devoted directly to the procurementof sufficient sustenance. In this particular historical context ritual maintenanceof cleared space is the method by which humans and their food become entangledin an obligate symbiotic relationship ( sensu Rindos, 1980).

Winterhalder and Goland (1997) predict that reliance on a series of highdensity, low rank resources would lead to local increases in population density.This increased population density could in turn lead to either intensified sub-sistence efforts or migration (or some combination of the two). Winterhalder and Golands model provides part of the logic for expecting the practice of ceremonial subsistence to spread. We predict this behavior spread either throughsimple biological fitness and expansion of populations of practitioners (Richersonet al., 2001:395; Rindos, 1980; Winterhalder and Goland, 1997:132, Figure 7.2)and/or through social learning within and among groups (e.g., Boyd andRicherson, 1985).

As the practitioners of this particular pattern of disturbance increased innumbers their neighbors would potentially see the benefits of their behavior andemulate them (i.e., social learning). This would result in the rapid spread of this method of interacting with the environment and any associated ideas andemic intentions. It is in this way that construction of earthworks and mounds(cultural elaboration) during the Woodland period becomes linked with fitness;not through wasteful expenditure of energy but through disguised productiveactivity that increases individual fitness relative to other contemporaneouspatterns of behavior (i.e., those not maintaining disturbed habitats).

The ceremonial subsistence model can potentially account for the widespreadpractice of mound construction and limited earthwork construction during theEarly Woodland period. If communities continued to interact in this way withtheir surroundings, then continued ritual activity, accumulation of new con-structions, and/or addition to existing works is required to maintain the level of disturbance necessary to retard succession in these areas. If populations did indeedincrease (as predicted by Winterhalder and Goland 1997), this would lead tothe expansion of these areas over centuries. This is congruent with the accretionalnature of Middle Woodland mound and earthwork complexes (e.g., Greber,1997). These expansions would continue to increase the abundance of these

food stuffs for exploitation by the populations building these earthworks.In good years the excess produce would not be needed and could be either leftto sow itself or be used in intergroup interaction through feasting and/or exchange.Whereas in bad years the buffer plots contained in the ceremonial centers wouldbe required for survival. In this way not only does this practice of disturbingthe area via ritual activity provide food stores for the lean season, it can be used



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to provide insurance during lean years via social storage. This is an especiallylikely scenario if each group has connections with groups in regions thatexperience non-correlated shortfalls (Kelly, 1995:Chapter 5; Winterhalder, 1986:Figure 6). This method of harvest variance management may, at least partially,explain the degree of exchange manifest during the Early and, especially, MiddleWoodland periods. A penchant for exotic material during the Middle Woodlandmay have been fueled by a situation characterized by low levels of environmentalvariability and low-levels of inter-group correlation in shortfalls and returns(Kelly, 1995:Figure 5-6). This could lead to a form of exchange involving goodsor services in addition to food (Winterhalder, 1986:387).

Our expectations for the pattern of Early and Middle Woodland period environ-mental variability stands in marked contrast to Dunnells (1996d, 1999; Dunnelland Greenlee, 1999) explanation for the Hopewell phenomenon. Dunnells modelrequires high levels of environmental variability to bring about elaboration; inour formulation, however, low levels of temporal variance are expected withmoderate to high levels of spatial variance (the spatial variance expectationis derived from Winterhalder, 1986). There is still the problem of specifyingquantitatively what the expected level of variation is for each model. Given thewide reaching nature of Hopewell exchange the scale of this spatial variabilitycould be quite large.

When considering temporal environmental variation it should be rememberedthat the focus of the mildly intensified productive effort are annual colonizingspecies, most of which we consider to be weeds today. As such they are relativelyinsensitive to fluctuations in weather and climate, reducing the potential for temporal variance in the subsistence system. Weed-like plant resources can act asa buffer against temporal variance in other resources, like nuts. Hickory, walnut,and acorn were an integral part of the diet of Archaic, and Woodland popula-tions of the Eastern Woodlands. While abundant in the region, these resourcesare known to produce in irregular cycles (Gardner, 1997:Figures 8.3-8.6;USDA, NRCS, 2008). In years with fewer nuts, the weedy plant resourceswould become crucial.

The End of Hopewell

Dunnell and Greenlee (1999) speculate that an increase in carrying capacity due to improved climatereduced variance in yields and alleviated the need todivert energy from reproductive efforts, bringing about the end of Hopewell

elaboration. According to evolutionary ecology models, if the climate hadimproved, the pressure to increase the amount of EAC crops in the area of settlement would have declined. Higher ranked, lower cost resources wouldhave become a greater and greater part of the diet until EAC crops were all butsqueezed out of the diet. Recent research suggests, however, that EAC cropsremained a dietary staple in the region into the Late Woodland period and beyond



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(Leone, 2007; Wymer, 1996, 1997; see also Nolan, 2009:Appendix B), well after the reduction in cultural elaboration and the suspected climate improvement.

We suspect there was a change internal to the system, such as a technologicalinnovation or a morphological change in one or a few of the crops, whichincreased the return rate for EAC cultigens. If the change was in the plant(s) andin the direction of increased dependence on human intervention, then indirectinvestment (via ceremonial activity) is no longer sufficient to maintain outputs.If the change is in technology and behavior, then efficient intensification wouldmake the community-level ceremonial aspect of subsistence superfluous. Witheither of these internal changes, energy for subsistence endeavors would need tobe redistributed most likely in the direction of increasingly intensive exploitationof whichever resources were the focus of change. If there was a change in theefficiency of seed crop exploitation (innovation or mutation), we would expectintensification of utilization of the affected resource(s) to the exclusion of other,now lower ranked, resources. Such a change would result in benefit to thoseexhibiting more direct investment in the growth, tending and harvesting of thisresource. Under these circumstances the efforts diverted to the communalbuffering plot (the ceremonial centers) would be abandoned in favor of moreefficient means of subsistence intensification. It should be noted that there islittle evidence of a change in environmental conditions at the time of the demiseof Hopewell (ca. AD 400) as called for by Dunnell and Greenlee (1999), even inthe sources they cite in support of their scenario.

An alternative explanation for the gradual cessation of ceremonial energyexpenditure is a long-term downturn in climatic conditions conducive to pro-duction. Bond et al. (2001) have documented quasi-periodic warm-cold cyclescorrelated with variation in solar productivity (insolation). Of particular interestis Bond event 1, a cold period of several hundred years duration (Yu et al.,2003:Figure 5) beginning around 2000 cal BP . This event is a long, gradualdecrease in temperatures that peaks sometime during the 5th century AD . TheBond cycles are argued to be large scale (hemispheric or global) climatic eventsrelated to irradiance (incoming solar radiation); however, the local consequencescan be variable. In North America, for example, the event is correlated with a dryspell in a western Canadian peat fen and in parts of Illinois and an increase inmoisture in southern Michigan (Booth and Jackson, 2003; Braun, 1987:168-169;Jackson and Booth, 2002; Yu et al., 2003). The take home message is that allof these trends begin around the same time (ca. AD 1-100) and extend beyondthe demise of the Hopewell phenomenon (i.e., past AD 400). If the lower

Great Lakes data are applicable to central and southern Ohio, then it wasincreasingly cold and wet during the Middle Woodland period. This wouldrepresent a gradual decline in the productivity of the environment throughoutthe period in which increasingly large areas were cleared for the earthwork/gardens. It is under these conditions that individual families or communitieswould have experienced initial success in their ritual/subsistence endeavors, but



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eventually more success in newly developed strategies for increasing subsistenceyield would be sufficient to overcome traditional emic reasons for carrying onthe cultural elaboration.

Under these circumstances, we predict decreased participation in the ritualaspects of subsistence at the communitylevel and increased attention to householdgardens. This strategy would require increased labor investment in gardeningand increasing levels of involvement in the plants life cycles. In addition to theprediction that labor investment in subsistence is intensified under times of stress,we base our predicted shift of energy to individual garden plots on Winterhalder and Golands (1997) model. They predict a shift of risk management strategiesto the household level as domesticates increase in rank (Winterhalder andGoland, 1997:141). Such an increase in rank and narrowing of the diet is predictedto happen under a coevolutionary model as domesticates compete with eachother in the agroecology (Rindos, 1980, 1984). Weeding, tilling, and other man-agement activities would become a regular part of the labor regime. Under conditions of changing climate and increasing human attention, EAC cultigensbegin to compete with each other, and the opportunity arises for stars in theagroecology. Rindos (1984) predicts that increasing investment in the agro-ecology will result in increasing specialization within the subsistence strategy.Those plants that respond most favorably to intensification will begin to receivemost of the human attention and become more prevalent in the diet and cor-responding archaeobotanical assemblages.

Braun (1987:169) reaches a similar conclusion. He points out that a changingclimate in Illinois may have selected against certain EAC plants, specificallymaygrass (adverse to cold) and marsh elder (preference for moist soils). He citesevidence of a cold/dry trend that starts north of Illinois around 2000 BP andcorresponds with the onset of his Trend 3 in the ceramic wall thickness data.The evidence from southern Michigan suggests regionally variable moistureregimes (Booth and Jackson, 2003), but otherwise the paleoclimatic evidencefor the Great Lakes region agrees with Brauns analysis. Thus, we might expectregionally specific changes in evenness of EAC seed quantities in archaeo-botanical assemblages; however, we still expect a change in human-plant inter-actions during this time. Brauns predicted subsistence strategy shift differs fromthe one predicted here, however. Where Braun (1987:169) expects a decline inthe contribution of cultigens to the diet under stress (actually stated as a slowingof intensification), we would predict an increase in subsistence intensificationduring these stressful times (Barlow, 2002).

Diet breadth models predict that during stressful times increasing amountsof energy will be invested in subsistence strategies (e.g., Barlow, 2002).Accordingly, if the environmental conditions deteriorated relative to needs for cultigen productivity, then intensification of subsistence effort is predicted. Thedirection we predict for this intensification is toward increasing investment inthe production of EAC cultigens and eventually maize.



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At first this intensification would be manifest as gradual expansion of earth-works and disturbed areas (during the Middle Woodland period), but at somepoint individual families would begin investing more effort directly in their ownproduction, curtailing their contribution to community wide efforts in theproduction of buffer crops (late Middle Woodland to early Late Woodland, circaAD 200-600). This leads to the development of a more specialized agroecology,and a more specialized subsistence strategy (at least with regards to cultigens).At this point it benefits our discussion to point out that Braun (1987:169) specu-lates that horticultural ecosystems changed in several ways during this time,possibly for several mutually reinforcing reasons leading up to the rapidincrease in maize use. This is an important point because of the nature of maize.The establishment of an agroecology and the transition into specialized domes-tication ( sensu Rindos, 1984) sets the stage for the intensification of maize pro-duction in the late Late Woodland.

Maize was a fully domesticated (morphological domesticate sensu Rindos,1984) import that was wholly dependent on humans for its survival. Maize hasbeen documented in the Eastern Woodlands during the Middle Woodland period(Hart, 1999:Table 1) in small quantities, but it was not a significant component of the diet at this time (Greenlee, 2002:Figure 24; Hart 1999:Figure 3; Winterhalder and Goland, 1997:145-146). Maize requires an established agroecology tothrive. The developments in horticultural ecosystems (a.k.a., agroecologysensu Rindos, 1980, 1984) during the transition from Middle to early LateWoodland period set the stage for the eventual development of maize-basedeconomies of the late Late Woodland and Late Prehistoric periods (ca. AD800-European contact). We want to emphasize, that from our perspective thetransition between Middle and Late Woodland periods is not a meaningful (real)event or period. We are simply using the terminological conventions currentlypredominant in the region.

Once a system of agricultural domestication ( sensu Rindos, 1980, 1984) wasestablished and maize had sufficient time to adapt to new climatic regimes of theEastern Woodlands (e.g., number of frost-free days) more successful varietiesof maize could become thriving members of the local agroecology (Hart, 1999).Once maize became suited to the local agroecology it would immediately beginto out compete its fellow inhabitants of the human-maintained environmentresulting in the sharp spike in apparent consumption ( d 13C values; Hart [1999:Figure 3]; though see Greenlee [2002:Figure 54]) and increased ubiquity inarchaeological features after AD 800.

As we have just illustrated, our model for the development of the Hopewellphenomenon couched within a historically contingent, coevolutionary trajectorycan reasonably lead to the trends documented for changes in subsistence after the decline of Hopewell in the Middle Ohio River Valley. More to the point,this scenario fits well with the climatic data currently available (Bond et al., 2001;Booth and Jackson, 2003; Hu et al., 2001; Jackson and Booth, 2002; Yu et al.,



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2003) indicating a long-term down-turn in environmental conditions conduciveto cultivation peaking in the mid-fifth century AD .

The strength of the ceremonial subsistence model is that it is built within ahistorically contingent scenario. The model does not assume that there is any-thing fundamentally (qualitatively) different about the Middle Woodland periodand recognizes that the developments during this time can be explained as acontinuation of long-term, local trends (i.e., going back to 6000 BP [the MiddleArchaic period] at least). We will now turn to a brief discussion of the (limited)extant evidence that can be used to evaluate this model of Middle Woodlandperiod cultural elaboration as a ceremonial subsistence strategy and severalempirical tests of this explanation that would discount this scenario and/or serveto draw distinctions between this explanation and the waste hypothesis.

Empirical Consequences

In this section we summarize the most significant empirical consequences for the ceremonial subsistence hypothesis. We evaluate these consequences againstthe limited available database (extremely limited in most cases) and the con-trasting empirical consequences of the waste hypothesis (see Table 2). We findthat the extant evidence fails to contrast with the expectations of the ceremonialsubsistence hypothesis, whereas in several instances the expectations of the wastehypothesis are contradicted. None of this analysis is sufficient to reject either hypothesis, but highlights the types of data generation that arenecessary to attemptto examine both explanations and distinguish the two alternatives empirically.

Floral Density and Distribution

This subsistence role for earthwork construction has one very critical empiricalconsequence related to the composition of paleofloral communities: the earth-works were replete with EAC crops. By way of a quick illustration of the foodproduction potential of these areas, they range in size from one or two smallenclosures (~30 m 30 m each; e.g., ~0.6 acre [0.253 ha] Reinhardt Complex[Nolan, 2009, 2010]) to dozens of mounds enclosed by expansive embankmentwalls (e.g., >130 acres [52.61 ha] enclosed at the Hopewell site [e.g., Weinberger,2009:Figure 2-1]). Smith (1992) estimates that, given the impressive nutritionalprofiles (1992:208) of the EAC domesticates a 1.2 acre (0.49 ha) garden couldfeed up to 10 people for up to 6 months. Phrased this way, the small ReinhardtComplex could feed the same number of people for 3 months. This is not an

insignificant buffer plot. The larger Hopewell complex could feed over 1000people for 6 months, and the larger Newark earthwork complex significantlymore. Of course these figures would depend on the entire surface area beingintensely farmed. This is not likely to have been the case. The point is that eventhe small ceremonial sites could serve as a substantial subsistence buffer andsupplement to the local carrying capacity.



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Table2.SummaryofEmpiricalConsequencesandEvidenceRequiredto

EvaluateAlternativeModels

Expectations

Waste

Cerem

onial

subsistence

Evidencepresentorneeded

Residentialstability

Ceremonialsiteslocatedatold

habitation/cemeterysites

Cultigenpollenatceremonialsites

Absenceofarborealpollenat

ceremonialsites

IncreaseEACubiquity

Intensifiedsubsistenceeffort

? ? ? ? ? No

Yes

Yes

Yes

Yes

Yes

Yes

Severalinstancesofhamlet(s)invicinityofceremonialsites(e.g.,

Pachecoetal.,2006,2009)

DatesspanningtheArchaic,andEarly/MiddleWoodland(e.g.,

Nolan,2010:Table6.1;RubyandLynott,2009:Table8-2)

PresenceinstratifiedcoresatFortAncient(McLauchlan,2003)

PresenceinstratifiedcoresatFortAncient(McLauchlan,2003)

Documentedathabitationsites(Wymer,1996,1997);insufficient

datafromceremonialsites

Increasingdisturbancetaxa(Wymer,1996,1997)andincreased

ubiquitycultigens(seeabove),increasinglylargecomplexesover

time(e.g.,Greber,1997)



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Lowharvestvariabilityandlow

intergroupharvestcorrelation

Highharvestvariability

Improvedclimate,increased

cultigenproductivityca.AD400

Decliningclimate,decreased

cultigenproductivityca.AD400

Diversificationofsettlement/

subsistencestrategiesca.AD200

Decreasedfertility(lessthan

maximalcarryingcapacity)

Increasedfertility/populationdensity

No

Yes

Yes N

o ? Yes N

o

Yes

No

No

Yes

Yes

No

Yes

Verylimitedavailabledataonfinescalevariability.

WarmermoisterclimateafterAD400(equivocal);endof

moundbuilding(present),anddecreasedimportanceofcultigens

(lacking;cf.Wymer,1996).Noreliablelocalempiricalinformation.

Intensifiedhouseholdgardens,abandonceremonialism(sameas

above);intensifieduseofsome

cultigens(i.e.,decreaseddiet

breadth;Wymer,1996);increasinglycold/moistclimatepeaking

ca.AD400(e.g.,Bondetal.,2001;Yuetal.,2001).Noreliablelocal

empiricalinformation.Needpollenanalysisatbasinsinother

ceremonialsites.

OriginofNewtownnucleatedsitesca.AD200(Dancey,1992).

Veryfewrelevantskeletalseries;possibilityofsteadyincreasein

fertilitythroughouttheWoodlandperiod(Bocquet-AppelandNaji,

2006;Buikstraetal.,1986).



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We must acknowledge being inspired here by the work of Kendra McLauchlan(2003). McLauchlan examined two sediment cores from depressions interpretedas artificial ponds within the embankments at Fort Ancient; one in the South Fortand one in the North Fort. The pollen in these cores is contained within a stratifiedcolumn from the accumulated sediments within the artificial ponds within thishilltop enclosure. The beginning of the record is set by the construction of theponds and anchored by radiocarbon dates from the core at ca. AD 1. McLauchlan,as is standard practice in pollen analysis and ecological reconstruction efforts,anchors her observed quantities of pollen types with a sequence of dates. There isa marked absence of arboreal pollen prior to AD 400, and a sharp transitionto predominantly arboreal pollen sometime after AD 200 (McLauchlan, 2003:Table 1, Figure 5).

McLauchlan reaches three major conclusions based on her analysis of thepollen in those two cores:

1. prehistoric agriculture was more extensive than previously thought(McLauchlan, 2003:564);

2. the nature and scale of human impacts on the landscape must be recon-sidered; and

3. that human impacts can be detected under appropriate conditions.

All of these are important with respect to the model presented here. The first andsecond conclusions clearly agree with the conclusions of Wymer (1996, 1997)about the nature of the subsistence economy at the Middle Woodland periodhabitation sites she examined. The third conclusion is of particular importancefor the evaluation of the ceremonial subsistence hypothesis.

If the ceremonial subsistence model is a valid explanation of Ohio Hopewellelaboration, then other mound and earthwork centers must have been kept inan arrested successional stage for their entire use-life. That is, most, if not all,earthworks must have evidence of similar pollen assemblages during the MiddleWoodland period (at least). Specifically, arboreal (or the local climax vegetation)pollen is expected to be a conspicuously minor component. Tree pollen is prolificand widely dispersed and is expected to dominate pollen diagrams in forestedareas. The absence of trees in the vicinity of earthworks (or even a significantlydecreased population) should be relatively easy to document in sediment coresnear earthworks. The key to McLauchlans (2003) third conclusion is that theright conditions must exist.

Any basin that receives water runoff from the surrounding area and remains

relatively undisturbed should serve as an acceptable sediment sampling site. If this is the case, there are myriad opportunities for testing this aspect of the modelat nearly every earthwork site in southern Ohio. If any of the borrow pits canbe demonstrated to have regularly received water runoff from the earthworksand vicinity, and especially if they were specifically constructed as ponds, asis suspected at Fort Ancient (McLauchlan, 2003:558), then they should bear



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evidence of prolonged deforestation and increased proportions of pollen fromdomesticated taxa (e.g., McLauchlan, 2003:Figures 5 and 7). However, hilltopenclosures may be better suited for this purpose. As tree pollen grains are smalland highly mobile they could easily contaminate terrace and floodplain sites.We expect that a pollen assemblage shift could still be detected at the boun-daries of Middle Woodland usage as was done for the Fort Ancient earthwork (McLauchlan, 2003:Figures 5 and 7). However, it may be more difficult todetermine the level of deforestation in lowland circumstances. Further pollensampling should be undertaken at Hopewell earthworks and enclosures toevaluate the expectations of the ceremonial subsistence model.

Prairie soils have been documented underlying earthworks at some Hopewellsites (Lepper, 1998; Nolan, 2010; Ruby, 1997). However, the presence of aprairie does not preclude the establishment of EAC plants. It should be remem-bered that the EAC plants have evolved to exploit the naturally disturbed habitatscreated by annual flooding, but they have broad environmental tolerance that permits them to colonize forest openings in a wide variety of settings (Gremillionet al., 2008:407; emphasis added). Therefore, there is no reason to think that apre-existing forest opening climax community precludes the operation of thismodel of earthwork construction. The pre-existing inhabitants (grasses) of theforest opening would still need to be disturbed and a deviation from the localsuccessional sequence maintained. However, earthworks constructed in areaswhere the climax groundcover is not arboreal may complicate detection of thepollen assemblage shift.

Subsistence Intensification and Changesin Settlement Pattern

The expectations for settlement patterns during the transition into the earlyLate Woodland (~ AD 200-400) are similar to those for Danceys (1992, 1996)model for the transition. Dancey (1992) detailed a model for shifts in settlementpatterns associated with changes in subsistence intensity, possibly associatedwith climatic deterioration. During his work on settlement patterns in centralOhio, Dancey noticed that a different pattern of settlement originated at a timeoverlapping with dwindling activity at the mound and earthwork centers. In1992, Dancey was not able to identify subsistence change as the likelycause of thistransition to village life; however, in his 1996 paper he postulates an intensifi-cation of the subsistence system as being responsible for this shift in settlement

strategy. Additionally, he notes that inception dates for Newtown early LateWoodland manifestations in southern Ohio range from AD 200400, indicatingsignificant overlap with Hopewell occupations (Dancey, 1996:399).

To clarify, we expect that, around AD 200 if not earlier (~200 years intoBond event 1), individual families or economic units began to aggregate awayfrom the dispersed hamlets associated with the earthworks in an attempt to



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intensify garden production due to declining return rates brought on by environ-mental deterioration (i.e., increasingly cold and wet conditions). This intensifi-cation and settlement shift is likely to take place first in less productive areasaway from the ceremonial centers and subsequently spread to the more productiveareas as the deterioration persists and/or the agricultural village communitiesbegan to experience a relative increase in reproductive success (Dancey, 1996:398, 399). If the newly aggregated and agricultural populations were successfulin their subsistence intensification this new strategy would spread.

We expect this to be the case for three reasons:

1. intensified production will increase food availability allowing higher population densities (relative to their less subsistence intensive neighbors)

(Richerson et al., 2001; Winterhalder and Goland, 1997);2. if the success of this system is perceptible by their neighbors, then it isexpected to spread via diffusion (social learning) into new areas (Boyd andRicherson, 1985); and finally

3. as Rindos (1980, 1984; Rindos and Johannessen, 1991) model of thedevelopment of agricultural subsistence strategies indicates, agriculturaladaptations are inherently unstable and this in itselfwill lead to the spread of this type of adaptation across the landscape (see also Richerson et al., 2001for a similar conclusion about the nature of the spread of agriculture).

If there was an increase in food stress during this time period we expect thatintensification of production would occur, resulting in a decrease in diet breadth;specifically we expect a shift in the proportional use of EAC plants. Someevidence for this type of a shift is provided by Wymer (1996). According toWymers interpretation of her data, in contrast to the Middle Woodland hamlets,early Late Woodland nucleated settlements were places where every availableresource seems to have been intensively utilized, including less desirable plantfoods (Wymer, 1996:42; emphasis added).

Wymer (1996) goes on to say that, while there is evidence of significant impacton the environment by Middle Woodland period communities, it is not to the samedegree evidenced by the few available early Late Woodland period nucleatedvillages. This is an important point with respect to the empirical consequences of the two alternative hypotheses under consideration. According to the scenariopresented by Dunnell and Greenlee (1999) subsistence intensification is notexpected. If the carrying capacity were to increase (for whatever reason) alowering in energy expenditure for food production would be expected. The

finding that early Late Woodland villagers were intensively utilizing all resourcesin the immediate environment does not point to any relief of food stress for thepost AD 400 occupants of the Middle Ohio River Valley (neither does the climateevidence reviewed above).

Elsewhere, Wymer (1997:158) asserts that the density and quantity of EACseeds in archaeobotanical assemblages is little changed from the Middle



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Woodland through the early Late Woodland. At the same time she argues that theMiddle Woodland people were farmers . This implies a high level of environ-mental disturbance and modification throughout the Woodland period (Wymer,1996:41, 1997:158; remember McLauchlans first and second conclusions).While the proportion of the EAC cultigens contribution to the diet is not known,Wymer (1996, 1997) and others (see Bellwood, 2005:174-179) argue for asubstantial contribution by these domesticated resources. A trend of high andrelatively stable reliance on domesticated resources and increasing levels of environmental disturbance during the early Late Woodland is congruent withthe model presented here and is a less good fit with the scenario proposed byDunnell and Greenlee (1999).

It is important to note that Wymers samples are small and geographicallycircumscribed. We do not mean to generalize her records to all Middle and LateWoodland sites. This is in fact, exactly the opposite of our approach. We aresimply highlighting that current interpretations of the limited relevant databaseagree with the expectations derived from our hypothesis and do not fit theexpectations derived from the alternative. This does not prove the case either way. A corollary point is that the empirical expectations derived from bothalternative hypotheses can and should be tested. The greatest use will come of our hypothesis if it generates new empirical work, even if the ceremonial sub-sistence hypothesis is rejected.

Under the waste hypothesis, conditions relevant to environmental produc-tivity (e.g., temperature, frost-free days, precipitation) were varying unpredictablyat the time scale of human experience during the Middle Woodland. In order todeal with these fluctuations, energy was diverted to non-reproductive behavior termedwaste.Theend of the Hopewell phenomenon wascatalyzed by ameliora-tion in the conditions relevant to environmental productivity. With the variation inthe subsistence system eliminated, wasteful behavior became inefficient and anyperson not participating in the elaborative behavior would experience increasedfitness relative to theirelaborating compatriots. The ceremonial subsistence modelis not dependent on an unpredictably fluctuating environment during the period of cultural elaboration and more importantly does not call upon that in a proximatecausal role. However, climatic deterioration is a probable forcing factor for thecessation of cultural elaboration during the Woodland period.

Fertility Rates

A final point of contrast between the empirical consequences of the ceremonialsubsistence and the waste hypotheses lies in the expectation for patterns infertility. A decrease in fertility coincident with increased cultural elaboration isat the core of Dunnells model. We would not expect a decrease throughout theperiod and especially not at the outset of mound and earthwork construction inthe region. Following the predictions of Winterhalder and Golands (1997) model,



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we would expect local population growth as the protodomesticates begin to beincluded in the optimal diet and especially as their density on the landscapeincreases through human modifications to the ecosystem. The relevant evidence islimited. Too few of the pertinent skeletal series have been examined, and thereis the problem of representativeness for those samples that are available, as mostcome from ceremonial sites and are not complete populations. Even more prob-lematic is that very few of the examined samples come from the Middle OhioRiver Valley.

Buikstra and colleagues (1986) have examined the relationship between thedevelopment of agriculture and fertility during the Woodland and Mississippianperiods in west-central Illinois. While their sample is small, they pick up a trendof increasing fertility from the Middle into the Late Woodland (Buikstra et al.,1986:Table 3, Figure 1). However, Buikstra et al. had no Early Woodland samplesand only two Middle Woodland samples and a total of eight. Additionally, thedifferences in the estimates of fertility only become significant when comparingthe earliest and the latest cemetery populations in their sample.

A more comprehensive data set has been compiled by Bocquet-Appel andNaji (2006). Bocquet-Appel and Naji (2006) pull together 60 cemeteries fromacross North America in their search for a universal demographic transition after the shift to a farming economy. There are sampling issues with this database(e.g., Eshed, 2006); however, they do examine 25 cemeteries from the Midwest(Bocquet-Appel and Naji, 2006:Figure 7; Chamberlain, 2006:Figure 1). Thesecemeteries span the time period ~4000 BC~AD 1400. These samples appear toshow a steady increase in population growth rate beginning at or before AD 1 andextending more or less steadily until ca. AD 1000. We must again urge caution ininterpreting these results; however, the available data do appear to contradictthe expectations of the waste hypothesis.

As Todd VanPool has pointed out to us, it is possible for a version of thewaste explanation to apply in a situation where absolute fertility is increasing(decreasing, or even staying the same as well). It is the relationship betweenmaximum potential carrying capacity and actual population size that the wastehypothesis addresses, not rate of reproduction per se. However, our discussionabout fertility rates applies equally well the proportional relationship betweenmaximum carrying capacity and actual population size. A strict contrast betweenthe two hypotheses would require much more detailed information about environ-mental capacity and Middle Woodland population levels than we currently haveaccess to. For now we are going to have to settle for the less direct approach

relevant to fertility rates.

SUMMARY AND CONCLUSION

The ceremonial subsistence model is reliant on Smiths (1987, 1989, 1995)scenario for the coevolution of the EAC plants-human relationship and its



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attendant requirements for a degree of permanence on the landscape. From thereit is expected that residentially stable communities were necessary for the main-tenance of central ceremonial-subsistence domestilocalities through the Early andinto the Middle Woodland periods. Finally, a gradual shift beginning aroundAD 200 toward nucleated settlements in habitats away from contemporarydispersed hamlets and associated vacant centers is expected, resulting in wide-spread intensified agricultural ( sensu Rindos, 1984) subsistence and nucleatedvillage settlements (Dancey, 1992, 1996).

While there is not enough evidence at present to fully test the model pre-sented here, the model cannot be rejected with the available evidence. Further-more, the few lines of evidence available for examination appear to contradictthe empirical implications of the waste hypothesis. Early Late Woodland periodsubsistence intensification (Wymer, 1996, 1997), the presence of pollen fromprobable cultigens within earthworks (McLauchlan, 2003), increasingly coldand wet climate during the Middle Woodland period (e.g., Bond et al., 2001;Booth and Jackson, 2003), and evidence of increasing fertility during the MiddleWoodland period (Bocquet-Appel and Naji, 2006; Buikstra et al., 1986) are inlinewith the empiricalexpectations of the ceremonialsubsistence modelpresentedhere and, to varying degrees, are at odds with the empirical expectations for thewaste explanation.

In concluding this discussion we call for heightened efforts to evaluate both of the models discussed in this article in various regions (but particularly the MiddleOhio River Valley). Both are based on a sound model of materialistic science(sensu Dunnell, 1996a, 1996b, 1996c, 1996d, 1996e) and as such have deducibleempirical consequences and can be rejected by detailed empirical work. Theareas of research that will be most useful in testing these explanations pertains tosettlement patterns before, during and after the Middle Woodland period; patternsin archaeobotanical and pollen assemblages from habitation and earthwork sites;detailed consideration of local climatic conditions throughout the span, and thespecific effects of those climatic conditions on subsistence; and, finally, anyevidence as to changes in fertility before, and during the Woodland period.

We hope that this article can serve as a call for an increase in theoretically-grounded, problem-oriented research into key issues surrounding the Hopewellclimax. Importantly we hope to highlight the importance of focusing investi -gation on both the preceding and following time periods in order to fully under-stand the context of this climax. We do not intend this article to necessarily bea rejection of the waste hypothesis in general or as applied to the Hopewell

case. Neither or both models may have explanatory power. The key is that bothmodels are derived from an explicit and consistent theoretical perspective andhave deducible empirical consequences. Both models need to be tightened withrespect to the nature of the specific expectations. As of now, the terms used in bothmodels are very coarse. However, having competing alternatives should drive usall to more critically evaluate these explanations and work toward more sufficient



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explanations. Finally, we hope that this article can serve as an example of how evolutionary archaeology and evolutionary ecology can be incorporatedinto a single explanatory narrative. It is increasingly recognized that these twoapproaches are not independent and antagonistic (Gremillion, 2006; Lipo andMadsen, 1998; Lyman and OBrien, 1998; see also Laland and Brown, 2002),but in fact need to be merged to produce complete explanations of the evolutionof cultural phenomena.

ACKNOWLEDGMENTS

Several people have helped us develop our thoughts and sharpen the expres-sion of our ideas. We both extend our appreciation to Kristen J. Gremillion for

reading multiple versions of this article and offering critical feedback that vastlyimproved our arguments. We also wish to thank William S. Dancey and JulieField for reading previous versions of this article and bluntly informing uswhen our arguments missed the mark. Dancey has also been an integral part inhelping both of us to think about these types of questions. Todd VanPool has alsooffered pointed, but constructive criticism on an earlier version of this article.Finally, we thank Reginald Byron for his feedback on the article. All of theseindividuals and the anonymous reviewers have helped us strengthen our argu-ments. Any remaining shortcomings, errors, and/or omissions in the article are,of course, our responsibility.

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using evolutionary archaeology and evolutionary ecology to explain cultural elaboration: the case of...

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