Using Evolutionary Archaeology and Evolutionary Ecology To Explain Cultural Elaboration: The Case of Middle Ohio Valley Woodland Period Ceremonial Subsistence

NORTH AMERICAN ARCHAEOLOGIST, Vol.
31(2) 119-154, 2010
USING EVOLUTIONARY ARCHAEOLOGY AND

EVOLUTIONARY ECOLOGY TO EXPLAIN CULTURAL
ELABORATION: THE CASE OF MIDDLE OHIO VALLEY
WOODLAND 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 Hopewell
phenomenon. 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 development
of the phenomenon being explained. Our model builds on Smith’s (1987)
coevolutionary scenario for the development of symbiosis between Eastern
Agricultural Complex (EAC) crops and the human populations exploiting
them. After presenting our model, we explore the empirical consequences
of ceremonial subsistence and the types of data that would be required to test
the model. In our discussion of the extant data we also contrast our scenario
with a competing evolutionary explanation for the Hopewell phenomenon:
the “waste” hypothesis. The limited available evidence is in line with the
empirical 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 annual
meeting of the Midwest Archaeological Conference at Notre Dame, Indiana.
119
Ó 2010, Baywood Publishing Co., Inc.

doi: 10.2190/NA.31.2.a
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120 / NOLAN AND HOWARD
degrees, at odds with the empirical expectations for the “waste” explanation.
Our ceremonial subsistence model offers a testable alternative that is not
falsified by the extant data. However, we recognize the problem is far from
solved. Our primary aim is to promote renewed theoretical discussion of the
issues raised herein and to encourage new problem-oriented research to
provide empirical evidence to test both explanations.
INTRODUCTION
Evolutionary archaeology and evolutionary ecology approaches to archaeology
are increasingly being recognized as complementary (e.g., Gremillion, 2006;
Lipo and Madsen, 1998; Lyman and O’Brien, 1998; see also Laland and Brown,
2002). We contribute to this growing trend by proposing a Darwinian evolutionary
model 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 an
alternative 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 the
available evidence, we identify an apparent lack-of-fit for the waste hypothesis
and describe the types of data required to fully evaluate both hypotheses. Due
to the current coarseness of both models and the limited available data, we are
unable 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 and
evolutionary ecology have been used to explain monumental architecture and
cultural elaboration such as that exhibited by the Ohio Hopewell. Each approach
has been successful at presenting plausible hypotheses for elaborative behavior
in various circumstances, and evolutionary archaeology has been applied
specifically to the Ohio Hopewell phenomenon (Dunnell, 1996d; Dunnell and
Greenlee, 1999). We feel that neither of these general explanations are a complete
explanation of Ohio Hopewell cultural elaboration. With a combination of
aspects of evolutionary archaeology and evolutionary ecology, we propose a
hypothesis that is plausible and empirically testable. We hope this article will
spur theoretical discussion and empirical evaluation of explanations such as the
ones discussed herein.
PREVIOUS EVOLUTIONARY MODELS OF

ELABORATIVE BEHAVIOR
Both evolutionary archaeologists and evolutionary ecologists note that cultural
elaboration is seemingly wasteful from a fitness standpoint, and both attempt to
demonstrate that it can be accounted for via individual-level fitness maximizing
CEREMONIAL SUBSISTENCE / 121
strategies. The “waste” hypothesis from evolutionary archaeology (e.g., Aranyosi,

1999; Dunnell and Greenlee, 1999; Hamilton, 1999; Kornbacher, 1999) and
“costly 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 provide
evolutionary explanations for seemingly inefficient behavior. Both hypotheses
draw upon Darwinian Theory and evolutionary explanations developed in
evolutionary 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, can
signal to females which one is more able to provide for offspring. The concept has
been employed convincingly to explain individual behaviors, such as material
display and risky hunting behavior (e.g., Bird and O’Connell, 2006). Although it
hasn’t been applied to the Hopewell situation, “costly signaling” theory has been
used to explain monument building in other regions. Neiman (1997) suggests
that Mayan elites were signaling their prowess to each other and prospective
followers through their erection of monumental architecture. In the case of the
Hopewell earthwork builders in the Middle Ohio Valley, there is no evidence
that any individual was ranked highly enough to be solely responsible for the
construction of the earthworks. Therefore, it would be unclear exactly who was
signaling. To date, no one has proposed a version of costly signaling as an
evolutionary explanation of the Hopewell phenomenon against which to evaluate
the evidence, so we focus our discussion and comparisons on the “waste”
hypothesis, which has been applied to the Hopewell situation.
The “waste” hypothesis was originally formulated by Dunnell in 1989 to
demonstrate the utility of an evolutionary archaeology approach (Dunnell, 1996d;
see also Dunnell, 1999), using the elaborative behavior of the Ohio Hopewell
as an illustrative example. Dunnell and Greenlee (1999) refined the argument
that mound building behavior diverts energy from reproductive behavior and
therefore is “wasteful,” apparently reducing the fitness of the individuals
involved. Dunnell (1999:246) argues that “waste” can be favored by “natural
selection acting in the usual fashion in somewhat unusual circumstances.” Within
(unusually) unpredictable environments, seemingly “wasteful” behavior can
actually benefit individual fitness by diverting energy into activities other than
production (and reproduction), thus reducing the numbers of offspring. This
keeps the population within the local carrying capacity during lean times (see
Dunnell, 1996d:Figure 4.2).
It is widely known in the biological sciences that organisms will divert energy
away from reproductive effort in temporally variable environments to increase
effective fitness (Dunnell, 1999:245; Madsen et al., 1999; see also Boyce and
Perrins, 1987; Seger and Brockman, 1987). In such environments it pays to
produce offspring below the organism’s maximum potential in order to raise
offspring of better competitive quality to the age of reproductive viability (Madsen
et al., 1999:Figure 3). The logic is that organisms will “sacrifice some expected
fitness so as to reduce their uncertainty, or variance, of fitness” (Seger and
Brockman, 1987:185). According to Seger and Brockman (1987) this is accom-
plished by maximizing the geometric-mean fitness (cf. arithmetic mean; see
also discussion in Madsen et al., 1999). This same logic may be applied to
human reproductive behavior. Madsen et al. (1999) used computer simulations to
demonstrate that, under the Darwinian theory of natural selection, seemingly
inefficient reproductive regimes will persist and in fact increase in frequency
under some circumstances. The simulations of Madsen et al. are re-illustrations
of the “geometric-mean principle” of Seger and Brockman (1987). However, in
the non-human cases that provide the theoretical basis for waste/bet-hedging
behavior, energy diverted from reproduction is not invested in elaborative
behavior. Excess energy can be reinvested in parenting effort, which makes
good Darwinian sense.
Alternatively, below optimum reproduction in fluctuating environments is
simply the best long-term “bet” to maximize individual lifetime reproductive
success (Boyce and Perrins, 1987). Variable environments select for reduced
production of offspring only, not for cultural elaboration. The proponents of the
“waste” hypothesis acknowledge that there is no necessary causal connection
(Madsen et al., 1999); however, they do not address how the linkage would
develop. For example, Dunnell and Greenlee (1999) focus only on how elabor-
ative behavior may have functioned and came to an end, with no attempt to
postulate how the linkage became established. If the “waste” hypothesis is to
be used to explain the origin (in addition to function, persistence, and cessation)
of elaborative behavior it seems necessary to address how the link between
fitness and elaboration becomes established. In other words, what is the historical
context within which the behavior(s) evolved?
Additional issues are raised by Madsen et al.’s (1999) reframing of Dunnell’s
original model. Dunnell’s 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 and
Brockman, 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 the
quality–vs.–quantity variety in their discussions of reproduction below optimum
levels 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 two
types of scenarios are also different. The trade-off (cost) hypothesis posits that
below maximum reproductive effort is selected for “when risk of adult mortality
increases with clutch size” (Boyce and Perrins, 1987:142). In contrast, the bet-
hedging type of reproductive restraint is predicted when there is increased
variability in survivorship of offspring (Boyce and Perrins, 1987:142; Seger and

Brockman, 1987). Therefore, bet-hedging and Dunnell’s original presentation of
“waste” are two theoretically possible evolutionary explanations for the regulation
of reproduction under different sets of circumstances.
Madsen et al. (1999) attempt to merge Seger and Brockman’s (1987) definitions
2.4 and 2.5 of bet-hedging with Dunnell’s original trade-off concept, but this is
problematic. Dunnell’s use of the energy trade-off establishes a relationship
between waste (non-reproductive use of energy) and long-term reproductive
success in varying environments. Madsen et al. (1999) attempt to rely mainly on
bet-hedging as defined by Seger and Brockman (1987) and observed by Boyce
and Perrins (1987) to explain increased elaborative behavior via a reduction in
fecundity, without establishing the connection between the two. The theory relied
on by Madsen et al. (1999) is capable of accounting for decreased fecundity
without cultural elaboration. Madsen et al. recognize this gap when they state
“that 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 addition
of elaboration would be necessary when the biological bet-hedging theory
is sufficient to explain reduced fecundity in temporally variable environments
without 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 are
simply re-illustrating the geometric-mean principle and make the unnecessary
assumption 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 is
empirical. More attention needs to be focused on the historically contingent set of
circumstances under which elaboration would become linked with a bet-hedging
type of regulation of production of offspring. Whether Dunnell’s version of
“waste” or Madsen et al.’s version bet-hedging/waste can explain a given set of
archaeological circumstances is an empirical rather than theoretical determination.
To clarify, the fact that Madsen et al. (1999) did not identify the empirical
historical circumstance that did or may have resulted in linking elaboration with
fecundity does not invalidate the theoretical argument in the case that such a link
can be established. Indeed, their paper was wholly theoretical and therefore could
not have identified a general historical case for the association between the two.
However, this absence does make the particular application difficult. For this
reason we attempt to trace the historical trajectory leading to elaboration.
The line of reasoning presented above does not weaken the general applicability
of either the trade-off or bet-hedging models to explain patterns in prehistory (not
limited to cultural elaboration). The overall logic of these explanations and their
connection to theory is sound. Dunnell’s (1996d) original intent of highlighting
the applicability of evolutionary theory to the archaeological record is important
to this discussion; however, his 1989 paper did not critically consider evidence in
support of his example. It simply illustrates how the logic of well established
evolutionary models could be applied to problematic cases in the archaeological
record; a lesson we see as crucial.
Bet-hedging (with or without elaboration) may very well explain many things
in the archaeological record. However, Madsen et al.’s (1999) specific application
to cultural elaboration does not seem to be sufficient. It is easy to imagine that
a few families not participating in elaborative behavior, but still observing below
optimal 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 not
plausible evolutionary explanations in some scenarios. We merely hope to illus-
trate by our theoretical analysis above and our presentation below that, given
the currently available data, the case for either or both of these hypotheses as
explanation for the Ohio Hopewell phenomenon is less than complete. First, the
link between elaboration and increased survivorship has not been established
by specific presentations of the waste hypothesis. Second, as we will show below,
the empirical record evidences several departures for the expectations derived
from the waste hypothesis.
If the development of earthwork construction in the Middle Ohio Valley
is to be accounted for by the action of natural selection (though there is
no necessary reason that this must be the case), then we argue that a
scenario that provides a direct contribution to fitness is more likely. It is
with the intent of exploring this possibility that we present the ceremonial sub-
sistence model.
CEREMONIAL SUBSISTENCE
In this section we lay out 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 the
development of the Hopewell climax (Middle Woodland) have their roots in the
Middle Archaic period (ca. 6000–3000 BC).
In our presentation we rely on the culture-historical terminology predominant
in the region (see Table 1). However, we should note that these terms are used in
various ways by various authors and in all cases they are largely common sense
terms and in need of theoretical justification for their definitions (e.g., Hart
and Brumbach, 2003). Most often the culture-historical terms are used to denote
Table 1. Generic Time Periods for Eastern North American Prehistorya
Period Subperiod Dates
Archaic Early 8000–6000 BC

Middle 6000–3000 BC
Late 3000–1000 BC
Woodland Early 1000 BC–AD 1000

Middle 1000–50 BC (Adena)
Late 50 BC–AD 400 (Hopewell)
Late Prehistoric AD 1000–1650 AD 400–1000

aGenerally associated cultural packages are shown in parentheses after the dates. Note
that there is no necessary association between the cultural package and the dates of the
period and that Adena and Hopewell are found to start at different times in different places.
units of space-time. This is the conceptually most prohibitive use and we avoid
that here. For ease of understanding, we use the terms Archaic, Woodland, and
their subdivisions as referents to blocks of time, not normative material packages
or cultural traditions (though the latter is their original meaning). We also,
sparingly and regrettably, use the terms Adena and Hopewell to refer to aggregate
packages of interpreted behavior. This usage, we fully acknowledge, and hope
becomes 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 Archaeology—employed
these terms is his argument for the waste hypothesis. These terms have wide
purchase in the region and are therefore retained here for efficiency of
communication at the expense of precision. We hope these choices do not cloud
out the nature of our argument.
As we proceed we will identify the types of evidence needed to evaluate
both the “waste” and ceremonial subsistence models and assess how each fit
the available data. Our goals in this article are to offer an alternative model
to compete with the waste model, and evaluate the veracity of both hypotheses
against 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 able
at 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. The
current data and interpretations discussed raise questions about the applic-
ability of the “waste” hypothesis and do not allow us to reject the ceremonial
subsistence hypothesis.
Historical Background for the Model

The basis for the ceremonial subsistence model is the development of the
Eastern Agricultural Complex (EAC), increased reliance upon food production
from the Middle Archaic period through the Middle Woodland period, and related
changes in settlement patterns (Smith, 1987, 1989, 1995; see also Braun, 1987).
The ceremonial subsistence model begins with Bruce Smith’s coevolutionary
scenario for the establishment of an association between human disturbed habitats
and abundance of edible plants.
Bruce Smith’s Floodplain Weed Theory “specifies how plant domestication and
cultivation grew out of behavioral responses [e.g., increasing residential stability
and 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 within
Gremillion’s (2002:Figure 22.3) zone of developed pre-maize agriculture (see
also Smith, 1989:Figure 1). We link the maintenance of disturbed habitats with the
practice of earthwork construction and posit that the continued evolution of EAC
plants takes place in association with, and perhaps partially within, the large
geometric 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 fruitful
avenue of research, recently producing numerous hypothetical, emic explanations
for various aspects of Hopewell cultural elaboration. Byers (2004) postulates that
the earthworks and their associated mortuary offerings are the result of world
renewal cults trying to maintain balance within the cosmos. Others have focused
on the geometric and astronomical aspects of the earthworks, positing that many
of the sites served as calendars (e.g., Romain, 2000). Carr (2006) suggests that,
in some cases, several earthworks in a region served differing, yet complementary
ceremonial functions, each important to the regional populations in their own
right. Lepper (2006) even hypothesizes that long distance pilgrimages figured
into 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 various
emic (proximate) explanations may be valid, and none would necessarily be
inconsistent with an overall evolutionary explanation (either bet-hedging/waste
or ceremonial subsistence). We are not concerned with emic aspects of the rituals
associated with earthwork construction. We are concerned with explanation at a
separate and not exclusive level.
The evolutionary approach asks different kinds of questions. Did the popu-
lations that participated in cultural elaboration, for whatever emic reason, receive
a 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 costly
alternatives were available? Could environmental, fitness-related changes have
led to the ultimate and abrupt abandonment of the practice across its entire
distribution? For our purposes, of more importance than the details of the
ceremony are the details of settlement and subsistence strategies.
There is much debate over the nature of the Early and Middle Woodland
period 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 pottery
first enters the record around 800 BC. Ceremonialism and trade became important
during the Early Woodland with the most visible signs of this found in the
more frequent construction of mounds and earthworks. Exotic materials become
regular and more prominent members of the artifact inventory. The new
ceremonial and exchange activities increased in frequency and quantity during
the Middle Woodland period with tons of soil and sediment moved to modify
the landscape and create large geometric enclosures, some containing multiple
mounds (e.g., Bernardini, 2004; Greber, 1997).
The most well known aspects of the Early and Middle Woodland periods are
the nature of the large earthworks that dotted much of the Eastern Woodlands
and 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 Middle
and 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 patterns
was sparked in the late 1980s and 1990s by Dancey and Pacheco (Dancey, 1991,
1992, 1998; Dancey and Pacheco, 1997a, 1997b; Pacheco, 1996, 1997). There are
essentially two views of Middle Woodland period settlement systems and there
is ongoing and vigorous debate between the two camps. Some researchers
contend that the Hopewell groups that constructed the massive earthworks were
“tribal societies . . . [that occupied] different sites . . . throughout the year” (Yerkes,
2009:119; see also Cowan, 2006; Yerkes, 2002, 2003). On the other end of
the debate, some researchers postulated that the Ohio Hopewell settlement
pattern conforms to the Vacant Center/Dispersed Hamlet model first put forth
by Olaf Prufer in 1965. The model holds that Hopewell family groups lived in
small, semi-permanent, homesteads dispersed around, but not within the “vacant”
ceremonial centers (Dancey and Pacheco, 1997b). “Vacant,” as it is used by

Dancey and Pacheco, and Prufer, does not mean devoid of human activity, or
archaeologically devoid of “habitation” debris. It simply means that there were
no permanent settlements within the ceremonial centers (Pacheco, 1996, 1997;
Prufer, 1996). This point seems to be continually lost on critics of the model, some
of whom consistently argue that the presence of any domestic debris at these
centers 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 settlement
pattern is difficult to assess with respect to what is expected for the model”
(Church and Ericksen, 1997:343). Pacheco et al. (2006, 2009) have recently
exposed two such hamlets within the catchment of the Liberty Earthworks near
Chillicothe, Ohio, clarifying to some extent the spatial relationship between
adjacent hamlets, and between hamlets and earthworks.
The elaborate behaviors associated with Hopewell dwindle and disappear at
the end of the Middle Woodland period around AD 400 (Dancey, 1996; Dunnell
and Greenlee, 1999). While the Late Woodland period in Ohio is little known
(Seeman and Dancey, 2000), the early Late Woodland (AD 400/500–700/800)
is generally characterized by large, sometimes fortified nucleated settlements
(Church, 1987; Church and Nass, 2002; Dancey, 1992, 1998; Seeman and
Dancey, 2000). The late Late Woodland (AD 700/800–1000) 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). Maize
becomes 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 residential

stability of Smith’s (1987, 1995) hypothesis for the origin of the EAC. Following
that trajectory forward in time, we expect relatively stable, long-term settlements
during the period of development of the ceremonial subsistence behavior (Early
and Middle Woodland). Using Bruce Smith’s 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 and
Pacheco, 1997a, 1997b; Pacheco, 1996a, 1996b; see also Prufer, 1996). While
weighing 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 to
particular habitation locations. We would point out that from an evolutionary
perspective, spatial and temporal variability in group residence patterns would

not be unexpected.
Inevitably, resources surrounding long-term settlements become depleted and
the habitation must be relocated. As settlements are relocated in previously
undisturbed 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 stages
leading to mature communities. As Gremillion et al. (2008:395) point out “[i]n
the humid temperate forests of eastern North America, the removal of mature
trees 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 seeds
and greens (e.g., Chenopoduim berlandieri [goosefoot], Helianthus annuus
[sunflower], Iva annua [marshelder/sumpweed]), temporarily increasing the
carrying capacity of the local environment.
If humans continued to disturb the soil in the vicinity of these older habitation
sites, the vegetation would remain in an artificially retarded successional stage
prolonging the elevation of the local carrying capacity. Any activity, regardless
of emic intent, that functioned to delay the onset of mature plant communities
would be favored in this way by natural selection. That is, if there is some reason
for people to continue to visit the disturbed areas associated with their earlier
habitations or cemeteries, then there would be an associated increase in the
quantity of these annual seeds and therefore local carrying capacity.
A possible proximate cause of the human population returning regularly is
to visit and/or inter their ancestors. Additionally, maintaining stands of nuts
(Gardner, 1997) and/or productive, ecotone hunting grounds could serve as
impetus for maintenance of forest openings (both of these would also directly
benefit fitness). Even small levels of human disturbance in the area would help
maintain the retarded successional state, resulting in maintenance of productive
patches of edible annual seeds and greens. Irrespective of the emic intent for
continued human presence and activity at these previously disturbed sites, any
activity that maintains an arrested state of succession will have the effect of
increasing the abundance of edible resources on the landscape, and will therefore
increase individual fitness and be favored by natural selection. Importantly,
the activities that are maintaining disturbed areas need not be intended by the
actors 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 is
relevant is whether or not performing that activity provides more resources than
not performing the activity, and whether or not performing that activity provides
more 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
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 population
density increases and a high risk subsistence system. Gradually, population
growth would apply stress to the subsistence system and increasing levels
of 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 associated
with early mound and earthwork construction prior to or at the advent of the
Woodland 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 areas
would have had the effect of increasing the productivity of the local environment.
With this mutual dependence established, any individual or family that invests
energy (for whatever proximate, emic intention) into maintaining areas of
disturbance on the landscape will increase their resource base. This would affect
not only the abundance of crops, but also, indirectly, the availability of game
animals in the vicinity. Deer are known to prefer open forest/forest edge situations
and 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 are
unlikely to frequent current habitation sites, the maintenance of open areas away
from living humans will have the effect of reducing search times for this highly
valued prey item and potentially buffering unsuccessful hunts with the bonus of
abundant plant harvests at the same 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 engages
in this type of maintained disturbance behavior. Relative to those who do not
revisit old disturbed areas this practice will result in more efficient returns on
energy expenditure, and, according to Winterhalder and Goland’s (1997) model,
increased local population.
The particular fashion by which the disturbed area is maintained will be a
historically contingent cultural behavior and may be inefficient as far as systems
of subsistence intensification are concerned; however, it does increase the density
of protodomesticates and the carrying capacity of the local catchment. The fact
that the EAC seeds are readily storable makes it likely that the advantage to
these 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 the
diet in the first place (Gremillion, 2004).
There are very obvious implications for the archaeobotanical assemblages

associated with this shift in subsistence. In fact, some of the expected changes
already have been documented in available data sets. For example, we expect an
increase through time in the ubiquity of EAC seeds and pollen at and near
earthwork and habitation sites (see McLauchlan, 2003; Wymer, 1996, 1997; and
discussion below). Braun (1987) identifies just such a pattern for the Illinois
Valley during the Early and Middle Woodland period associated with trends in
ceramic vessel wall thickness.
Woodland Period Monumental Architecture
With the well established symbiosis and the selective advantage conferred
on agents of disturbance, any set of behaviors that perpetuates this disturbance is
beneficial. 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 to
the incipient agroecology (sensu Rindos, 1980, 1984) created by these ceremonial
sites and other open areas, selection will continue to favor those individuals
(humans) that perpetuate these open areas. By the Early Woodland period, we
know that groups of people began to erect monuments associated with mortuary
behavior. Whatever the emic reason, the disturbances associated with this activity
would function to maintain an early stage of succession (e.g., see pollen diagrams
in McLauchlan, 2003). The mortuary and ceremonial practices of the preceding
period were sufficient to maintain modest increases in domesticate productivity,
but the addition of moderate to large scale movement of earth increases the level
of 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 (perhaps
even one). Eventually the practitioners of this variety of disturbance were more
successful at maintaining higher yields over long periods of time. Thus the
practice 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 wouldn’t the prehistoric incipient
horticulturalists simply plant gardens? Why build earthworks and maintain
ceremonial sites to indirectly increase subsistence yields?” We expect that house-
holds did maintain independent gardens. If people simply intended to maximize
their long-term subsistence yield in the most direct fashion they would likely
skip the community-level ceremonial aspects of subsistence production (indeed,
we expect this eventually happened). However, individual intentions have little
role to play in long-term evolutionary narratives. Emic intentions can serve in
a proximate causal role for part of the historical sequence. The incipient farmers
of the Ohio Valley did not intend to become agriculturalists (see especially
Rindos’ [1980:769-770] reply to commentators; also contributions to Price and
Gebauer, 1995).
Evolution is historical and opportunistic. Natural selection acts on extant

variability; it does not forge it de novo. In our scenario, the ritual behaviors
associated with maintenance of cleared space and eventually massive, regular
movements of dirt would increase the return rate of “foraging” efforts thereby
decreasing the level of time and energy devoted directly to the procurement
of sufficient sustenance. In this particular historical context ritual maintenance
of cleared space is the method by which humans and their food become entangled
in an obligate symbiotic relationship (sensu Rindos, 1980).
Winterhalder and Goland (1997) predict that reliance on a series of high
density, 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 Goland’s model provides part of the logic for expecting the practice of
ceremonial subsistence to spread. We predict this behavior spread either through
simple biological fitness and expansion of populations of practitioners (Richerson
et 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 and
Richerson, 1985).
As the practitioners of this particular pattern of disturbance increased in
numbers their neighbors would potentially see the benefits of their behavior and
emulate them (i.e., social learning). This would result in the rapid spread of
this method of interacting with the environment and any associated ideas and
emic 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 productive
activity that increases individual fitness relative to other contemporaneous
patterns of behavior (i.e., those not maintaining disturbed habitats).
The ceremonial subsistence model can potentially account for the widespread
practice of mound construction and limited earthwork construction during the
Early Woodland period. If communities continued to interact in this way with
their 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 indeed
increase (as predicted by Winterhalder and Goland 1997), this would lead to
the expansion of these areas over centuries. This is congruent with the accretional
nature 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 left
to 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 would
be required for survival. In this way not only does this practice of disturbing
the area via ritual activity provide food stores for the lean season, it can be used
to provide insurance during lean years via social storage. This is an especially
likely scenario if each group has connections with groups in regions that
experience 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, Middle
Woodland periods. A penchant for exotic material during the Middle Woodland
may have been fueled by a situation characterized by low levels of environmental
variability 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 goods
or 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 Dunnell’s (1996d, 1999; Dunnell
and Greenlee, 1999) explanation for the Hopewell phenomenon. Dunnell’s model
requires high levels of environmental variability to bring about elaboration; in
our formulation, however, low levels of temporal variance are expected with
moderate to high levels of spatial variance (the spatial variance expectation
is derived from Winterhalder, 1986). There is still the problem of specifying
quantitatively what the expected level of variation is for each model. Given the
wide reaching nature of Hopewell exchange the scale of this spatial variability
could be quite large.
When considering temporal environmental variation it should be remembered
that the focus of the mildly intensified productive effort are annual colonizing
species, most of which we consider to be weeds today. As such they are relatively
insensitive to fluctuations in weather and climate, reducing the potential for
temporal variance in the subsistence system. Weed-like plant resources can act as
a 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 resources
are known to produce in irregular cycles (Gardner, 1997:Figures 8.3-8.6;
USDA, NRCS, 2008). In years with fewer nuts, the weedy plant resources
would become crucial.
The End of Hopewell
Dunnell and Greenlee (1999) speculate that an increase in carrying capacity—

due to improved climate—reduced variance in yields and alleviated the need to
divert energy from reproductive efforts, bringing about the end of Hopewell
elaboration. According to evolutionary ecology models, if the climate had
improved, the pressure to increase the amount of EAC crops in the area of
settlement would have declined. Higher ranked, lower cost resources would
have become a greater and greater part of the diet until EAC crops were all but
squeezed out of the diet. Recent research suggests, however, that EAC crops
remained a dietary staple in the region into the Late Woodland period and beyond
(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 technological
innovation or a morphological change in one or a few of the crops, which
increased the return rate for EAC cultigens. If the change was in the plant(s) and
in the direction of increased dependence on human intervention, then indirect
investment (via ceremonial activity) is no longer sufficient to maintain outputs.
If the change is in technology and behavior, then efficient intensification would
make the community-level ceremonial aspect of subsistence superfluous. With
either of these internal changes, energy for subsistence endeavors would need to
be redistributed most likely in the direction of increasingly intensive exploitation
of whichever resources were the focus of change. If there was a change in the
efficiency of seed crop exploitation (innovation or mutation), we would expect
intensification of utilization of the affected resource(s) to the exclusion of other,
now lower ranked, resources. Such a change would result in benefit to those
exhibiting more direct investment in the growth, tending and harvesting of this
resource. Under these circumstances the efforts diverted to the communal
buffering plot (the ceremonial centers) would be abandoned in favor of more
efficient means of subsistence intensification. It should be noted that there is
little evidence of a change in environmental conditions at the time of the “demise”
of Hopewell (ca. AD 400) as called for by Dunnell and Greenlee (1999), even in
the sources they cite in support of their scenario.
An alternative explanation for the gradual cessation of ceremonial energy
expenditure is a long-term downturn in climatic conditions conducive to pro-
duction. Bond et al. (2001) have documented quasi-periodic warm-cold cycles
correlated with variation in solar productivity (insolation). Of particular interest
is 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, gradual
decrease in temperatures that peaks sometime during the 5th century AD. The
Bond cycles are argued to be large scale (hemispheric or global) climatic events
related to irradiance (incoming solar radiation); however, the local consequences
can be variable. In North America, for example, the event is correlated with a dry
spell in a western Canadian peat fen and in parts of Illinois and an increase in
moisture 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 all
of these trends begin around the same time (ca. AD 1-100) and extend beyond
the “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 was
increasingly cold and wet during the Middle Woodland period. This would
represent a gradual decline in the productivity of the environment throughout
the period in which increasingly large areas were cleared for the earthwork/
gardens. It is under these conditions that individual families or communities
would have experienced initial success in their ritual/subsistence endeavors, but
eventually more success in newly developed strategies for increasing subsistence

yield would be sufficient to overcome traditional emic reasons for carrying on
the cultural elaboration.
Under these circumstances, we predict decreased participation in the ritual
aspects of subsistence at the community level and increased attention to household
gardens. This strategy would require increased labor investment in gardening
and increasing levels of involvement in the plants’ life cycles. In addition to the
prediction 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 Goland’s (1997) model. They predict a shift of risk management strategies
to the household level as domesticates increase in rank (Winterhalder and
Goland, 1997:141). Such an increase in rank and narrowing of the diet is predicted
to happen under a coevolutionary model as domesticates compete with each
other 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 cultigens
begin to compete with each other, and the opportunity arises for “stars” in the
agroecology. 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 receive
most 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 changing
climate in Illinois may have selected against certain EAC plants, specifically
maygrass (adverse to cold) and marsh elder (preference for moist soils). He cites
evidence of a cold/dry trend that starts north of Illinois around 2000 BP and
corresponds with the onset of his Trend 3 in the ceramic wall thickness data.
The evidence from southern Michigan suggests regionally variable moisture
regimes (Booth and Jackson, 2003), but otherwise the paleoclimatic evidence
for the Great Lakes region agrees with Braun’s analysis. Thus, we might expect
regionally 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. Braun’s predicted subsistence strategy shift differs from
the one predicted here, however. Where Braun (1987:169) expects a decline in
the contribution of cultigens to the diet under stress (actually stated as a slowing
of intensification), we would predict an increase in subsistence intensification
during these stressful times (Barlow, 2002).
Diet breadth models predict that during stressful times increasing amounts
of 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. The
direction we predict for this intensification is toward increasing investment in
the production of EAC cultigens and eventually maize.
At first this intensification would be manifest as gradual expansion of earth-

works and disturbed areas (during the Middle Woodland period), but at some
point individual families would begin investing more effort directly in their own
production, curtailing their contribution to community wide efforts in the
production of buffer crops (late Middle Woodland to early Late Woodland, circa
AD 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 “rapid
increase 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 has
been 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 to
thrive. The developments in “horticultural ecosystems” (a.k.a., agroecology
sensu Rindos, 1980, 1984) during the transition from Middle to early Late
Woodland period set the stage for the eventual development of maize-based
economies of the late Late Woodland and Late Prehistoric periods (ca. AD
800-European contact). We want to emphasize, that from our perspective the
“transition” between Middle and Late Woodland periods is not a meaningful (real)
event or period. We are simply using the terminological conventions currently
predominant in the region.
Once a system of agricultural domestication (sensu Rindos, 1980, 1984) was
established and maize had sufficient time to adapt to new climatic regimes of the
Eastern Woodlands (e.g., number of frost-free days) more successful varieties
of maize could become thriving members of the local agroecology (Hart, 1999).
Once maize became suited to the local agroecology it would immediately begin
to out compete its fellow inhabitants of the human-maintained environment
resulting in the sharp spike in apparent consumption (d13C values; Hart [1999:
Figure 3]; though see Greenlee [2002:Figure 54]) and increased ubiquity in
archaeological features after AD 800.
As we have just illustrated, our model for the development of the “Hopewell
phenomenon” couched within a historically contingent, coevolutionary trajectory
can 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.,
2003) indicating a long-term down-turn in environmental conditions conducive

to cultivation peaking in the mid-fifth century AD.
The strength of the ceremonial subsistence model is that it is built within a
historically contingent scenario. The model does not assume that there is any-
thing fundamentally (qualitatively) different about the Middle Woodland period
and recognizes that the developments during this time can be explained as a
continuation of long-term, local trends (i.e., going back to 6000 BP [the Middle
Archaic 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 Woodland
period cultural elaboration as a ceremonial subsistence strategy and several
empirical tests of this explanation that would discount this scenario and/or serve
to 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 against
the limited available database (extremely limited in most cases) and the con-
trasting empirical consequences of the waste hypothesis (see Table 2). We find
that the extant evidence fails to contrast with the expectations of the ceremonial
subsistence hypothesis, whereas in several instances the expectations of the waste
hypothesis are contradicted. None of this analysis is sufficient to reject either
hypothesis, but highlights the types of data generation that are necessary to attempt
to examine both explanations and distinguish the two alternatives empirically.
Floral Density and Distribution
This subsistence role for earthwork construction has one very critical empirical
consequence related to the composition of paleofloral communities: the earth-
works were replete with EAC crops. By way of a quick illustration of the food
production potential of these areas, they range in size from one or two small
enclosures (~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 embankment
walls (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 nutritional
profiles” (1992:208) of the EAC domesticates a 1.2 acre (0.49 ha) garden could
feed up to 10 people for up to 6 months. Phrased this way, the small Reinhardt
Complex could feed the same number of people for 3 months. This is not an
insignificant buffer plot. The larger Hopewell complex could feed over 1000
people for 6 months, and the larger Newark earthwork complex significantly
more. Of course these figures would depend on the entire surface area being
intensely farmed. This is not likely to have been the case. The point is that even
the small ceremonial sites could serve as a substantial subsistence buffer and
supplement to the local carrying capacity.
Table 2. Summary of Empirical Consequences and Evidence Required to
Evaluate Alternative Models
Ceremonial
Expectations Waste subsistence Evidence present or needed
Residential stability ? Yes Several instances of hamlet(s) in vicinity of ceremonial sites (e.g.,
Pacheco et al., 2006, 2009)
Ceremonial sites located at old ? Yes Dates spanning the Archaic, and Early/Middle Woodland (e.g.,
habitation/cemetery sites Nolan, 2010:Table 6.1; Ruby and Lynott, 2009:Table 8-2)
Cultigen pollen at ceremonial sites ? Yes Presence in stratified cores at Fort Ancient (McLauchlan, 2003)
Absence of arboreal pollen at ? Yes Presence in stratified cores at Fort Ancient (McLauchlan, 2003)
ceremonial sites
Increase EAC ubiquity ? Yes Documented at habitation sites (Wymer, 1996, 1997); insufficient
data from ceremonial sites
Intensified subsistence effort No Yes Increasing disturbance taxa (Wymer, 1996, 1997) and increased
ubiquity cultigens (see above), increasingly large complexes over
time (e.g., Greber, 1997)
Low harvest variability and low No Yes
intergroup harvest correlation
Very limited available data on fine scale variability.
High harvest variability Yes No
Improved climate, increased Yes No Warmer moister climate after AD 400 (equivocal); end of
cultigen productivity ca. AD 400 moundbuilding (present), and decreased importance of cultigens
(lacking; cf. Wymer, 1996). No reliable local empirical information.
Declining climate, decreased No Yes Intensified household gardens, abandon ceremonialism (same as
cultigen productivity ca. AD 400 above); intensified use of some cultigens (i.e., decreased diet
breadth; Wymer, 1996); increasingly cold/moist climate peaking
ca. AD 400 (e.g., Bond et al., 2001; Yu et al., 2001). No reliable local
empirical information. Need pollen analysis at basins in other
ceremonial sites.
Diversification of settlement/ ? Yes Origin of Newtown nucleated sites ca. AD 200 (Dancey, 1992).
subsistence strategies ca. AD 200
Decreased fertility (less than Yes No

maximal carrying capacity) Very few relevant skeletal series; possibility of steady increase in
fertility throughout the Woodland period (Bocquet-Appel and Naji,
Increased fertility/population density No Yes 2006; Buikstra et al., 1986).
CEREMONIAL SUBSISTENCE
/ 139
We must acknowledge being inspired here by the work of Kendra McLauchlan

(2003). McLauchlan examined two sediment cores from depressions interpreted
as artificial ponds within the embankments at Fort Ancient; one in the South Fort
and one in the North Fort. The pollen in these cores is contained within a stratified
column from the accumulated sediments within the artificial ponds within this
hilltop enclosure. The beginning of the record is set by the construction of the
ponds 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 is
a marked absence of arboreal pollen prior to AD 400, and a sharp transition
to predominantly arboreal pollen sometime after AD 200 (McLauchlan, 2003:
Table 1, Figure 5).
McLauchlan reaches three major conclusions based on her analysis of the
pollen 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 and
second conclusions clearly agree with the conclusions of Wymer (1996, 1997)
about the nature of the subsistence economy at the Middle Woodland period
habitation sites she examined. The third conclusion is of particular importance
for the evaluation of the ceremonial subsistence hypothesis.
If the ceremonial subsistence model is a valid explanation of Ohio Hopewell
elaboration, then other mound and earthwork centers must have been kept in
an arrested successional stage for their entire use-life. That is, most, if not all,
earthworks must have evidence of similar pollen assemblages during the Middle
Woodland period (at least). Specifically, arboreal (or the local climax vegetation)
pollen is expected to be a conspicuously minor component. Tree pollen is prolific
and widely dispersed and is expected to dominate pollen diagrams in forested
areas. The absence of trees in the vicinity of earthworks (or even a significantly
decreased population) should be relatively easy to document in sediment cores
near earthworks. The key to McLauchlan’s (2003) third conclusion is that the
right 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 model
at nearly every earthwork site in southern Ohio. If any of the borrow pits can
be demonstrated to have regularly received water runoff from the earthworks
and vicinity, and especially if they were specifically constructed as ponds, as
is suspected at Fort Ancient (McLauchlan, 2003:558), then they should bear
evidence of prolonged deforestation and increased proportions of pollen from

domesticated taxa (e.g., McLauchlan, 2003:Figures 5 and 7). However, hilltop
enclosures may be better suited for this purpose. As tree pollen grains are small
and 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 to
determine the level of deforestation in lowland circumstances. Further pollen
sampling should be undertaken at Hopewell earthworks and enclosures to
evaluate the expectations of the ceremonial subsistence model.
Prairie soils have been documented underlying earthworks at some Hopewell
sites (Lepper, 1998; Nolan, 2010; Ruby, 1997). However, the presence of a
prairie does not preclude the establishment of EAC plants. It should be remem-
bered that the EAC plants “have evolved to exploit the naturally disturbed habitats
created by annual flooding, but they have broad environmental tolerance that
permits them to colonize forest openings in a wide variety of settings” (Gremillion
et al., 2008:407; emphasis added). Therefore, there is no reason to think that a
pre-existing forest opening climax community precludes the operation of this
model of earthwork construction. The pre-existing inhabitants (grasses) of the
forest opening would still need to be disturbed and a deviation from the local
successional sequence maintained. However, earthworks constructed in areas
where the climax groundcover is not arboreal may complicate detection of the
pollen assemblage shift.
Subsistence Intensification and Changes

in Settlement Pattern
The expectations for settlement patterns during the transition into the early
Late Woodland (~AD 200-400) are similar to those for Dancey’s (1992, 1996)
model for the transition. Dancey (1992) detailed a model for shifts in settlement
patterns associated with changes in subsistence intensity, possibly associated
with climatic deterioration. During his work on settlement patterns in central
Ohio, Dancey noticed that a different pattern of settlement originated at a time
overlapping with dwindling activity at the mound and earthwork centers. In
1992, Dancey was not able to identify subsistence change as the likely cause of this
transition 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 Late
Woodland manifestations in southern Ohio range from AD 200–400, indicating
significant overlap with Hopewell occupations (Dancey, 1996:399).
To clarify, we expect that, around AD 200 if not earlier (~200 years into
Bond event 1), individual families or economic units began to aggregate away
from the dispersed hamlets associated with the earthworks in an attempt to
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 areas
away from the ceremonial centers and subsequently spread to the more productive
areas as the deterioration persists and/or the agricultural village communities
began to experience a relative increase in reproductive success (Dancey, 1996:
398, 399). If the newly aggregated and agricultural populations were successful
in 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 is
expected to spread via diffusion (social learning) into new areas (Boyd and
Richerson, 1985); and finally
3. as Rindos’ (1980, 1984; Rindos and Johannessen, 1991) model of the
development of agricultural subsistence strategies indicates, agricultural
adaptations are inherently unstable and this in itself will lead to the spread of
this type of adaptation across the landscape (see also Richerson et al., 2001
for 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 that
intensification of production would occur, resulting in a decrease in diet breadth;
specifically we expect a shift in the proportional use of EAC plants. Some
evidence for this type of a shift is provided by Wymer (1996). According to
Wymer’s interpretation of her data, in contrast to the Middle Woodland hamlets,
early Late Woodland nucleated settlements were places “where every available
resource seems to have been intensively utilized, including less desirable plant
foods” (Wymer, 1996:42; emphasis added).
Wymer (1996) goes on to say that, while there is evidence of significant impact
on the environment by Middle Woodland period communities, it is not to the same
degree evidenced by the few available early Late Woodland period nucleated
villages. This is an important point with respect to the empirical consequences of
the two alternative hypotheses under consideration. According to the scenario
presented by Dunnell and Greenlee (1999) subsistence intensification is not
expected. If the carrying capacity were to increase (for whatever reason) a
lowering in energy expenditure for food production would be expected. The
finding that early Late Woodland villagers were intensively utilizing all resources
in the immediate environment does not point to any relief of food stress for the
post AD 400 occupants of the Middle Ohio River Valley (neither does the climate
evidence reviewed above).
Elsewhere, Wymer (1997:158) asserts that the density and quantity of EAC
seeds in archaeobotanical assemblages is little changed from the Middle
Woodland through the early Late Woodland. At the same time she argues that the
Middle 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 McLauchlan’s 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 a
substantial contribution by these domesticated resources. A trend of high and
relatively stable reliance on domesticated resources and increasing levels of
environmental disturbance during the early Late Woodland is congruent with
the model presented here and is a less good fit with the scenario proposed by
Dunnell and Greenlee (1999).
It is important to note that Wymer’s samples are small and geographically
circumscribed. We do not mean to generalize her records to all Middle and Late
Woodland sites. This is in fact, exactly the opposite of our approach. We are
simply highlighting that current interpretations of the limited relevant database
agree with the expectations derived from our hypothesis and do not fit the
expectations derived from the alternative. This does not “prove” the case either
way. A corollary point is that the empirical expectations derived from both
alternative 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 unpredictably
at the time scale of human experience during the Middle Woodland. In order to
deal with these fluctuations, energy was diverted to non-reproductive behavior
termed “waste.” The end of the Hopewell phenomenon was catalyzed by ameliora-
tion in the conditions relevant to environmental productivity. With the variation in
the subsistence system eliminated, “wasteful” behavior became inefficient and any
person not participating in the elaborative behavior would experience increased
fitness relative to their elaborating compatriots. The ceremonial subsistence model
is not dependent on an unpredictably fluctuating environment during the period of
cultural elaboration and more importantly does not call upon that in a proximate
causal role. However, climatic deterioration is a probable forcing factor for the
cessation of cultural elaboration during the Woodland period.
Fertility Rates
A final point of contrast between the empirical consequences of the ceremonial

subsistence and the “waste” hypotheses lies in the expectation for patterns in
fertility. A decrease in fertility coincident with increased cultural elaboration is
at the core of Dunnell’s model. We would not expect a decrease throughout the
period and especially not at the outset of mound and earthwork construction in
the region. Following the predictions of Winterhalder and Goland’s (1997) model,
we would expect local population growth as the protodomesticates begin to be

included in the optimal diet and especially as their density on the landscape
increases through human modifications to the ecosystem. The relevant evidence is
limited. Too few of the pertinent skeletal series have been examined, and there
is the problem of representativeness for those samples that are available, as most
come from ceremonial sites and are not complete populations. Even more prob-
lematic is that very few of the examined samples come from the Middle Ohio
River Valley.
Buikstra and colleagues (1986) have examined the relationship between the
development of agriculture and fertility during the Woodland and Mississippian
periods in west-central Illinois. While their sample is small, they pick up a trend
of 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 samples
and only two Middle Woodland samples and a total of eight. Additionally, the
differences in the estimates of fertility only become significant when comparing
the earliest and the latest cemetery populations in their sample.
A more comprehensive data set has been compiled by Bocquet-Appel and
Naji (2006). Bocquet-Appel and Naji (2006) pull together 60 cemeteries from
across 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). These
cemeteries span the time period ~4000 BC–~AD 1400. These samples appear to
show a steady increase in population growth rate beginning at or before AD 1 and
extending more or less steadily until ca. AD 1000. We must again urge caution in
interpreting these results; however, the available data do appear to contradict
the expectations of the “waste” hypothesis.
As Todd VanPool has pointed out to us, it is possible for a version of the
waste explanation to apply in a situation where absolute fertility is increasing
(decreasing, or even staying the same as well). It is the relationship between
maximum potential carrying capacity and actual population size that the waste
hypothesis addresses, not rate of reproduction per se. However, our discussion
about fertility rates applies equally well the proportional relationship between
maximum carrying capacity and actual population size. A strict contrast between
the two hypotheses would require much more detailed information about environ-
mental capacity and Middle Woodland population levels than we currently have
access 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 Smith’s (1987, 1989, 1995)

scenario for the coevolution of the EAC plants-human relationship and its
attendant requirements for a degree of permanence on the landscape. From there

it is expected that residentially stable communities were necessary for the main-
tenance of central ceremonial-subsistence domestilocalities through the Early and
into the Middle Woodland periods. Finally, a gradual shift beginning around
AD 200 toward nucleated settlements in habitats away from contemporary
dispersed hamlets and associated “vacant” centers is expected, resulting in wide-
spread intensified agricultural (sensu Rindos, 1984) subsistence and nucleated
village 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 contradict
the empirical implications of the “waste” hypothesis. Early Late Woodland period
subsistence intensification (Wymer, 1996, 1997), the presence of pollen from
probable cultigens within earthworks (McLauchlan, 2003), increasingly cold
and wet climate during the Middle Woodland period (e.g., Bond et al., 2001;
Booth and Jackson, 2003), and evidence of increasing fertility during the Middle
Woodland period (Bocquet-Appel and Naji, 2006; Buikstra et al., 1986) are in
line with the empirical expectations of the ceremonial subsistence model presented
here and, to varying degrees, are at odds with the empirical expectations for the
“waste” 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 Middle
Ohio River Valley). Both are based on a sound model of materialistic science
(sensu Dunnell, 1996a, 1996b, 1996c, 1996d, 1996e) and as such have deducible
empirical consequences and can be rejected by detailed empirical work. The
areas of research that will be most useful in testing these explanations pertains to
settlement patterns before, during and after the Middle Woodland period; patterns
in archaeobotanical and pollen assemblages from habitation and earthwork sites;
detailed consideration of local climatic conditions throughout the span, and the
specific effects of those climatic conditions on subsistence; and, finally, any
evidence 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 Hopewell
“climax.” 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 be
a 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 both
models are derived from an explicit and consistent theoretical perspective and
have deducible empirical consequences. Both models need to be tightened with
respect to the nature of the specific expectations. As of now, the terms used in both
models are very coarse. However, having competing alternatives should drive us
all to more critically evaluate these explanations and work toward more sufficient
explanations. Finally, we hope that this article can serve as an example of

how evolutionary archaeology and evolutionary ecology can be incorporated
into a single explanatory narrative. It is increasingly recognized that these two
approaches are not independent and antagonistic (Gremillion, 2006; Lipo and
Madsen, 1998; Lyman and O’Brien, 1998; see also Laland and Brown, 2002),
but in fact need to be merged to produce complete explanations of the evolution
of 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 vastly
improved our arguments. We also wish to thank William S. Dancey and Julie
Field for reading previous versions of this article and bluntly informing us
when our arguments missed the mark. Dancey has also been an integral part in
helping both of us to think about these types of questions. Todd VanPool has also
offered pointed, but constructive criticism on an earlier version of this article.
Finally, we thank Reginald Byron for his feedback on the article. All of these
individuals 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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accepted for publication.
Direct reprint requests to:

Kevin Nolan
Department of Anthropology
The Ohio State University
4034 Smith Laboratory
174 W. 18th Avenue
Columbus, OH 43210-1106
e-mail: kcnolan49@gmail.com

Using Evolutionary Archaeology and Evolutionary Ecology To Explain Cultural Elaboration: The Case of Middle Ohio Valley Woodland Period Ceremonial Subsistence

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Using Evolutionary Archaeology and Evolutionary Ecology To Explain Cultural Elaboration: The Case of Middle Ohio Valley Woodland Period Ceremonial Subsistence

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NORTH AMERICAN ARCHAEOLOGIST, Vol.

31(2) 119-154, 2010

USING EVOLUTIONARY ARCHAEOLOGY AND

We propose a Darwinian evolutionary model for the development and dis-

Ó 2010, Baywood Publishing Co., Inc.

PREVIOUS EVOLUTIONARY MODELS OF

strategies. The “waste” hypothesis from evolutionary archaeology (e.g., Aranyosi,

variability in survivorship of offspring (Boyce and Perrins, 1987:142; Seger and

Table 1. Generic Time Periods for Eastern North American Prehistorya

Period Subperiod Dates

Archaic Early 8000–6000 BC

Woodland Early 1000 BC–AD 1000

Late Prehistoric AD 1000–1650 AD 400–1000

Historical Background for the Model

The Ohio Hopewell and the Rise and Fall of Elaboration

ceremonial centers (Dancey and Pacheco, 1997b). “Vacant,” as it is used by

The ceremonial subsistence model begins with the increasing residential

perspective, spatial and temporal variability in group residence patterns would

There are very obvious implications for the archaeobotanical assemblages

Woodland Period Monumental Architecture

Evolution is historical and opportunistic. Natural selection acts on extant

The End of Hopewell

Dunnell and Greenlee (1999) speculate that an increase in carrying capacity—

eventually more success in newly developed strategies for increasing subsistence

At first this intensification would be manifest as gradual expansion of earth-

2003) indicating a long-term down-turn in environmental conditions conducive

Floral Density and Distribution

Expectations Waste subsistence Evidence present or needed

Decreased fertility (less than Yes No

We must acknowledge being inspired here by the work of Kendra McLauchlan

evidence of prolonged deforestation and increased proportions of pollen from

Subsistence Intensification and Changes

intensify garden production due to declining return rates brought on by environ-

A final point of contrast between the empirical consequences of the ceremonial

we would expect local population growth as the protodomesticates begin to be

SUMMARY AND CONCLUSION

The ceremonial subsistence model is reliant on Smith’s (1987, 1989, 1995)

attendant requirements for a degree of permanence on the landscape. From there

explanations. Finally, we hope that this article can serve as an example of

BIRD, DOUGLAS, and JAMES F. O’CONNELL

CHURCH, FLORA, and JOHN P. NASS

1996e Archaeology and Evolutionary Science, in Evolutionary Archaeology:

LYMAN, R. LEE, and MICHAEL J. O’BRIEN

1997 Ohio Middle Woodland Intracommunity Settlement Variability: A Case

WINTERHALDER, BRUCE, and CAROL GOLAND

Direct reprint requests to:

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