Particularism and the Retreat from Theory in the Archaeology of Agricultural Origins Kristen J

Home , Historical particularism

SPECIAL FEATURE: PERSPECTIVE PERSPECTIVE SPECIAL FEATURE: Particularism and the retreat from theory in the archaeology of agricultural origins Kristen J. Gremilliona,1, Loukas Bartonb, and Dolores R. Pipernoc,d aDepartment of Anthropology, Ohio State University, Columbus, OH 43210; bDepartment of Anthropology, University of Pittsburgh, Pittsburgh, PA 15260; cDepartment of Anthropology, Smithsonian National Museum of Natural History, Washington, DC 20013-7012; and dSmithsonian Tropical Research Institute, Apartado Postal 0843-03092, Panama, Republic of Panama

Edited by James O’Connell, University of Utah, Salt Lake City, UT, and approved October 28, 2013 (received for review June 26, 2013)

The introduction of new analytic methods and expansion of research into previously untapped regions have greatly increased the scale and resolution of data relevant to the origins of agriculture (OA). As a result, the recognition of varied historical pathways to agriculture and the continuum of management strategies have complicated the search for general explanations for the transition to food production. In this environment, higher-level theoretical frameworks are sometimes rejected on the grounds that they force conclusions that are incompatible with real-world variability. Some of those who take this position argue instead that OA should be explained in terms of local and historically contingent factors.This retreat from theory in favor of particularism is based on the faulty beliefs that complex phenomena such as agricultural origins demand equally complex explanations and that explanation is possible in the absence of theoretically based assumptions. The same scholars who are suspicious of generalization are reluctant to embrace evolutionary approaches to human behavior on the grounds that they are ahistorical, overly simplistic, and dismissive of agency and intent. We argue that these criticisms are misplaced and explain why a coherent theory of human behavior that acknowledges its evolutionary history is essential to advancing understanding of OA. Continued progress depends on the integration of human behavior and culture into the emerging synthesis of evolutionary developmental biology that informs contemporary research into plant and animal domestication.

evolutionary theory | behavioral ecology

Over the last decade there has been a major This trend, coupled with the critical stance In arguing for the systematic use of theory expansion of knowledge regarding the timing toward hypothetico-deductive science cur- as a tool for the advancement of knowledge, and socioecological context of plant domesti- rently influential in archaeological thought we are not making the claim that evolution- cation and emerging agricultural systems. (15), has motivated many archaeologists to arytheoryistheonlypossiblerealmfrom This wealth of data is due in large part to favor interpretive modes of explanation. In which to choose. However, neo-Darwinism methodological innovations (e.g., in genetics origins of agriculture (OA) research, one has withstood repeated challenges to its core and paleogenomics, in the analysis of plant result of this changing climate has been a elements, having earned status as an over- micro- and macroremains and biological res- proliferation of historical narratives that arching framework for explaining the di- idues, and in the physical and biogeochemical highlight local events and processes and versity of life. For this reason, evolutionary analyses of anthropogenic sediments), reex- downplay general principles. Although we theory is central to understanding the root cavations of some important archaeological recognize the value of empirical rigor, we causes of human behavior and indeed culture sites, and the expansion of archaeological re- argue that the progress of OA research is itself (16). An even stronger case for the evo- search into regions whose record of agricul- significantly hampered by the denial of lutionary perspective on human behavior can tural origins has been until recently poorly explanations derived from a systematic be made now that the distinctive features of known [such as New Guinea (1, 2), lowland program of theoretically driven hypothesis cultural transmission and evolution are be- areas of Mesoamerica and northern South testing. We also reject assertions that evo- coming more fully incorporated into the syn- America (3–7), and northern and southern lutionary theory is teleological, needlessly thesis (e.g., refs. 17–20). China (8–13)]. These research activities have reductionist, ahistorical, and therefore irrel- We divide our discussion of these issues enriched both the scale and resolution of the evant to understanding human behavior. into three sections. The first elaborates upon data relevant to agricultural origins world- Without a coherent theory of human behav- some of the weaknesses of particularism in wide. One result of this welcome enhance- ior, arguments about agricultural origins OA research. Next, we sharpen the focus ment of the empirical record is wider must rely on assumptions that are often in- to respond to critiques of evolutionary acknowledgment of the variability in the his- tuitive, unstated, and largely implicit. Cou- approaches to human behavior [particularly torical pathways taken by emerging food pro- pled with the increasing emphasis on the research program of human behavioral duction systems across space and time. The human exceptionalism, the erosion of scien- ecology (HBE) and one of its tools, optimal dichotomy between foraging and food pro- tific method is troubling because it creates foraging theory (OFT)]. We argue that recent duction has been discarded in favor of a con- a rift between researchers in biology and ar- calls to abandon HBE or OFT are misguided, tinuum of landscape, plant, and animal chaeology whose collaboration is essential to management strategies that sometimes resist advancing understanding of OA. To address Author contributions: K.J.G., L.B., and D.R.P. wrote the paper. fi classi cation. However, for some scholars (2, these weaknesses, we contend that evolution- The authors declare no conflict of interest. 14), the richly detailed records of change ary theory (broadly construed to include cul- This article is a PNAS Direct Submission. seem to have dampened the appeal of general tural as well as biological processes) must 1To whom correspondence should be addressed. E-mail: gremillion. explanations for the transition to agriculture. play a central role in OA research. [email protected].

www.pnas.org/cgi/doi/10.1073/pnas.1308938110 PNAS | April 29, 2014 | vol. 111 | no. 17 | 6171–6177 Downloaded by guest on September 28, 2021 and we explain why the particularist alter- (23). The result is a rejection of both uni- historical accounts of OA (see, for exam- natives might provide more explanatory versal causation and general principles in ple, refs. 25 and 29-32). However, to the depth and power when situated within favor of “frameworks of explanation that extent that they purport to engage in ex- a program of behavioral ecology. Finally, pay close and careful attention to existing planation, such accounts often rely on we assess the prospects for a synthetic relevant archaeological information, that inductions that are expected to stand on evolutionary research program for the are scaled at the regional level, and that the strength of available evidence alone. study of domestication and agriculture. focus on the complex interplay of a range of Although we recognize that the prioritiza- different environmental and social pre- tion of data is often intentional and explicitly Limitations of Particularism in OA conditions, prompts, and factors of various stated (e.g., ref. 33), we disagree that it is Research and the Need for Theory kinds” (ref. 23, p. 681). a good idea to abandon the iterative process The research trend that we critique in this We agree that no single cause explains all of hypothesis testing, revision, and retesting paper shares several characteristics with the instances of transition from foraging to food that drives incremental advances in scientific approach labeled historical particularism by production (25–27); this observation is true understanding. historians of anthropology (21). Most closely but trivial. An explanation that claims uni- associated with Franz Boas and his students, versal validity loses its universality when Particularistic Explanations Lack a General historical particularism emphasized the col- exceptions are found to exist. However, its Theory for the Evolution of Human Behavior lection and compilation of ethnographic data remaining components—variables, processes, Yet Rely on a General Theory of Evolution without the distorting effects of theory. Boas and assumptions— often remain relevant at for All Other Organisms. Thereislittledis- and his adherents, weary of unsubstantiated a more modest level of generality. The fatal agreement that change in the frequency of theorizing unsupported by rigorous field- flaw in the prime mover arguments is not attributes in populations of humans, animals work, viewed theory as an obstacle to scien- that they rely on overarching laws or and plants under domestication is inherently tific investigation. Generalizations about assumptions, but rather that they are ap- coevolutionary. Understanding this process culture were suspect because exceptions to plied at an inappropriate scale. Following requires a general theory that links change in them could always be found. Any reliable the same logic, the optimization assumptions one population to change in another. Cer- generalizations would be derived inductively used heuristically in many HBE models tainly, humans possess a unique mode of from large bodies of data. Similar recom- should not be confused with a claim that inheritance and an unusual degree of be- mendations and concerns have been voiced economic optimization explains all tran- havioral flexibility. However, cultural in- in the OA literature (14, 22–24), with the sitions to agriculture (ref. 23, p. 682). heritance and the mechanics of human important difference that current expressions We agree that multiple broad-scale and learning that enable it are constrained (at of particularism, rather than disavowing the- local factors are involved in any agricultural least, in the long run) by processes that ory, embed it in arguments without explicit transition (25, 26). However, this is a com- constrain the evolution of other organisms. acknowledgment or relegate it to a minor role monsense observation about the world, not Far from being an obstacle to evolutionary that prioritizes empirically based inference. a template for explanation (ref. 23, p. 681). In analysis, human uniqueness offers one of the fi We argue that the retreat from theory and fact, complex phenomena such as OA must most important justi cations for using it. the embrace of particularism are not only be simplified to be understood. A theoretical By not interpreting human behavior as unwarranted but also counterproductive. framework is needed to guide this process to a product of natural selection, particularistic Particularistic approaches that reject gener- ensure that explanations are not fashioned approaches miss the opportunity to explore alization fail to treat inductive inferences as from a hodgepodge of factors and variables commonalities in human behavior that en- hypotheses to be tested and substitute appeals selected at the discretion of the researcher. able comparative analysis of agricultural ori- to agency for a consistent theory of human For this reason, we advocate an approach to gins in starkly different contexts. Nothing behavior. These strategies are problematic for understanding OA that examines theoretical about an evolutionary perspective suggests reasons we detail below. assumptions before interpreting data. Such that natural selection generates a pre- assumptions always exist, acknowledged or determined template for human behavior, Particularistic Approaches Reject Gen- not. Brought to the forefront, they serve to nor does it claim to explain everything that eral Principles in the Mistaken Belief anchor observations to bodies of knowledge humans do. Human social learning has its That They Imply Universal Causes and that have withstood repeated testing. Such own distinctive processes that influence pat- Ignore History. In the current climate, it is high-level bodies of general theory inform terns of behavior at the population level. not surprising that some OA researchers call middle- and lower-level theories that in turn However, human abilities (including the for a more particularistic approach to un- generate testable hypotheses (16, 28). The capacity for communication, learning, and derstanding why human groups became in- explanatory relevance of broadly applicable choice—all components of cultural involved in plant cultivation and animal principles does not conflict with the obser- heritance) are products of natural selection, husbandry. Among the reasons given for vation that domestication of any plant or and human culture is a function of those rejecting generalizations about causality is animal is a particular, local, and historically abilities (16, 34–36). Because agriculture has that contradictory cases can be answered contingent process. If it did, evolutionary been adopted (or inherited) by nearly all only by adding qualifiers (ref. 2, p. S380). theory could not be said to explain the di- human groups and because it was invented Some investigators conflate universal ex- versity of life without implying that all evo- separately by many, the process will be un- planations (i.e., “prime movers” like envi- lutionary lineages follow the same path. derstood best by looking at change in the ronmental change, population pressure, or To be clear, we applaud the attention being history of human adaptation as a function of divine inspiration), which imply that all paid to accurate reconstructions of the tra- attributes shared by all humans, rather than transitions to agriculture conform to the jectory and context of agricultural transitions looking at novel attributes unique to a few. same causal sequence, with the principles at regional and local scales. Detail-oriented Because they lack a general theory for and assumptions of HBE, which do not analysis of observations and data yield rich human behavior, particularistic approaches

6172 | www.pnas.org/cgi/doi/10.1073/pnas.1308938110 Gremillion et al. Downloaded by guest on September 28, 2021 rely on diverse and often conflicted principles analysis than the simple arithmetic required The Study of Evolution Is a Historical PERSPECTIVE to account for the decision-making behavior to find optimal solutions to foraging prob- Science. Evolutionary theory is not covering SPECIAL FEATURE: of human agents. In some cases, they retain lems (for example) and is even farther re- law science (ref. 23, p. 681). It is a scientific the rational-actor optimization logic of HBE moved from the practical heuristics (e.g., approach to history (47) that views change as (e.g., that people will act in their own best rules of thumb, tacit or explicit learning, ra- both cause and consequence of the immedi- interest with regards to survival-related utility); tional calculation, etc.) that people actually ate and long-term fitness implications of in other cases, they rely on creative or arbitrary use to make those decisions. By contrast, inherited and acquired variation. Although combinations of a “bounded rationality” simple agency in foraging models is one of focused on the processes of change, nothing (sensu refs. 37 and 38) drawn opportunis- individual decision making, but does not about these processes, or the analysis of them, tically from ethnographic or contemporary imply (and does not require) long-term is deterministic, teleological, or scripted. The observations; still others seem to gather teleological self-determination. context and consequences of one case might principles of behavior from social theory be completely different from the next (i.e., The Evolutionary Approach Is a Robust, (39, 40). Such principles are seldom made both historical and particular), but the foun- Flexible, and Practical Avenue for an explicit; in some cases they are revealed by dations of evolutionary analysis are unifying. fi references to unde ned or vaguely concep- Incrementally Better Understanding of Atthesametime,itisflexible enough to deal “ tualized properties or goals [e.g., resource- Human Behavior and Cultural Change with the hyper-variability of human behavior, ”“ fi rich, intensi cation of resource manipula- Whereas evolutionary biologists seldom have which, for a variety of evolutionarily sound ”“ tion, rich array of plant and animal spe- to confront the question of human excep- reasons, evolves and diversifies rapidly (48). ”“ ”“ ” cies, enrichment, stable and sustainable, tionalism, the issue is a central one in an- In part, this flexibility comes from the ap- “ ” stress, etc. (23)]. Such terms betray em- thropology and archaeology. The application plication of a wide range of models that bedded assumptions about the ecological of evolutionary theory to human behavior isolate specific aspects of behavioral and situations that humans prefer and pre- has been criticized for excessive adaptation- evolutionary phenomena. sumably strive for. Consistent theory that ism, reductionism, positivism, and material- illustrates how and why these organizing ism (e.g., refs. 43 and 44). These criticisms Simple Models of Complex Phenomena fi principles emerge in the rst place is have been taken up by many advocates of Are Useful Even When They Fail. Recent essential for explaining the outcomes of interpretive archaeology, who question the reactions to the use of OFT’s simplifying diverse motivations in the distant past, exclusive claim of science to the production models in hunter–gatherer and OA research particularly when these outcomes are visible of useful knowledge (many examples appear (14, 49, 50) take their cue from older critiques today only in the aggregate records of ar- in ref. 45; see also ref. 24). in biology and ecology (e.g., refs. 51 and 52) chaeology and population biology. Some archaeologists who do embrace but differ on the issue of culture. For many Researchers who reject general theories of evolutionary concepts in their OA research social science scholars, models that isolate human behavior and cultural evolution as remain uncomfortable with their application specific aspects of individual behavior in a overly deterministic sometimes invoke human to human behavior. Particularly heavy criti- ceteris paribus world (53) are impractical agency as an alternative causal mechanism. cism has been leveled at evolutionary ecology because (i) the ceteris paribus world doesn’t Agency is a contested term in archaeology, (EE), defined as the “study of evolution and exist, and (ii) individuals perform in a and the OA literature seems to have adopted adaptive design in ecological context” (ref. 46, complex, indivisible milieu that can be un- one sense of it to the exclusion of others. p. 3), especially the subdiscipline of HBE and derstood only in its holistic totality. For Advocates use the common reading of its optimal foraging models. These critiques many, the practice of methodological iso- agency (refs. 15 and 41, p. 67) to illustrate target either the basic assumptions of EE lation and analytical simplification has how individual choice and creativity explain (many of which are shared by other evolu- “ cultural change, including the transition to tionary approaches) or elements of HBE or reduced human actors to disemboweled fi humanswhonolongerhavecultural agriculture(ref.29,p.44).Althoughweac- OFT speci cally (14, 22, 23). Although the ” knowledge the role of human creativity and objections are numerous, they commonly anchors (ref. 54, p. 59). They complain that innovation in OA, we are not convinced that revolve around the following themes: (i) evolutionary models isolate individuals from appeals to agency have much explanatory explanations based on evolution by natural populations and remove both from the thing power, simply because they sidestep the selection are “covering law” explanations that binds them, which is culture. problem of identifying the evolutionary basis that fail to account for the particular his- Reservations about heuristic models are of the beliefs and preferences that guide hu- torical circumstances of their social and welcome (indeed these reservations and man action (42). In contrast, HBE generates biological contexts; (ii) simple models are the resulting critiques may lead to better hypotheses about why humans behave the inappropriate for complex phenomena; models); outright rejection of them is not. way they do and evaluates whether or not (iii) the use of multiple simplifying models Unfortunately, and in many cases, rejection these hypotheses about agency explain the is compromised when one or more models of simple, heuristic models is based on an material remains of those behaviors. We fail to explain or predict the phenomena at idealized (but often unstated and un- agree that human agency is a critical com- hand; (iv) models do not explain human supported) view of how the world works and ponent of evolutionary change, but we do not subsistence (or any other) behavior; (v)ex- on the complaint that too many people take see it as having an equally important role in planatory, predictive models deny human the modeled predictions as fact, with or the scientific explanation of change. agency and are therefore dehumanizing; and without empirical backing (14). Such cri- Another problem with explanation by (vi) some models do not account for envi- tiques mistakenly conclude that the optimi- agency is that it often implies that decision ronmental change and are therefore in- zation assumptions used heuristically in makers have complete information about the appropriate for historical or evolutionary many HBE models constitute a claim that long-term consequences of their actions. This analysis. We find these critiques unwarranted optimal resource use explains all transitions feat would require far more involved data and explain our reasoning below. to agriculture (ref. 23, p. 682).

Gremillion et al. PNAS | April 29, 2014 | vol. 111 | no. 17 | 6173 Downloaded by guest on September 28, 2021 The use of simple models (like those used ant to OA research. Aggregate patterns of on small-sized dietary items, occurred as in OFT) does not imply that the predictions individual behavior are what we see in the a result of climate- and/or human-driven of the models must be true. All things being archives of the past. Therefore, the study of factors, depending on the region involved. equal (in the ceteris paribus world), the models the past requires evaluation of models Examples include terminal Pleistocene fau- should explain the behavior of organisms against data aggregated over different scales. nal extinctions and vegetational change foraging optimally to maximize reproductive TheDBMisonemodelthatmakesthis (Neotropics) and increasing human de- success. In the absence of data, the modeled procedure possible. It is not the only model, mographic pressure leading to resource predictions are simply that, and if the and there is no reason to expect it to explain depression (southwest Asia). It is impor- assumptions are correct, the predictions all data on all scales of time or space. How- tant to reiterate (88) that ethnographic simply ought to be true. When data do exist ever, in combination with related models, it is datasets on foraging efficiency from around but do not fit the model, either all things are a very powerful analytical tool. the world support the ranking of general not equal (e.g., the assumptions are in- categories of resources such as small and correct), the model is inappropriately ap- A Challenge to One Model Does Not large game according to their energetic return plied (e.g., the scale or resolution of the Negate the Others. Onecritiqueofthe rates (net energy obtained per unit time behaviors does not match those of the evolutionary program is that it shifts arbi- spent). Although exceptions occur, most resulting data and/or the currency of the trarily between models that isolate and ad- plant foods and small and medium-sized model is inappropriate), or the organism dress different aspects of phenomena staged mammals are ranked lower than most large ’ simply doesn t conform to expectation. at different scales of time, space, or number game, and nuts and seeds are often the low- fi Observations of the nal type can also be (e.g., ref. 67). The observation is fair, but the est-ranked of all (80, 88). It would be coun- productive by leading to further hypothe- critique is not. Indeed, what makes EE so “ terproductive for investigations of cultural sis testing built on related versions of EE versatile is that it draws from a family of developments through time, such as agricul- fi models” to address the tradeoffs between developed speci cally to explain the pro- tural origins, to ignore those associations. generality, realism, and precision inherent to liferation and persistence of suboptimal or The collective archaeological data clearly maladaptive behavior (17, 55–58). any approach to scientificinquiry(68–70). underline how dietary diversification and The criticism that HBE models sometimes Theimportantpointhereisthatwedrawon intensification of plant use by foragers and fail to yield accurate predictions when tested models rooted in a unified body of theory, incipient farmers probably occurred in an misses the point that the exploration of the even if some of the models fail to explain the environmental milieu of shifting energetic model’s vulnerability can itself be a source of existing data, or if the models explain the return rates. Arguments that DBM pre- insight. Models are productive means for data on different scales or levels of analysis. dictions for the BSR are frequently invali- “eliminating problematic answers and iden- ” Models of EE Often Generate Accurate dated by the archaeological data (14) are tifying and pursuing more promising ones fi (ref. 59, p. 171). In other words, failure is an Predictions of Human Subsistence Be- dif cult to reconcile with the results of these option. For example, when empirical tests of havior, Including OA. As previously noted, studies. As a number of scholars have the diet breadth model (DBM) failed to show OFTisoneclassofmodelswithareasonable pointed out, the DBM is particularly suited a good fit of data to model predictions for purchase in the explanatory realm, and one for studying major directional changes in agricultural transitions in the eastern United that has provided a number of insights for subsistence through time because of its ability States, the findings led to a better grasp of the biologists and archaeologists. A review of the to make robust, qualitative predictions of variables and tradeoffs (such as travel and past few years of biological literature finds prey choice and dietary diversity. Using the processing costs) that most influenced sub- persistent employment of OFT models, with model does not require that every past or sistence strategies (e.g., refs. 60 and 61). empirical data often showing good fits with present resource ranking set be in perfect Piperno and Pearsall (62) also discussed how their predictions in a variety of organisms order with respect to the general category of OFT models, whose predictions conformed (e.g., refs. 71–74). OFT applications in hu- resource, package size, and the energetic to Neotropical empirical data on the emer- man research also continue apace with return rates of individual items. gence of food production, were unlikely valuable insights on varied topics (e.g., refs. to address subsequent agricultural in- 75–78). Ecological Models Accommodate Human tensification satisfactorily, which evoked A particularly important and well-studied Agency. Rejection of models like the DBM other explanations. Others (63–65) have in- example of subsistence change associated in favor of explanation by agency echoes vestigated the failure of simple foraging with OA and one that has been a focus of a long-standing debate in evolutionary bi- – models such as the DBM to capture im- OFTapplicationsisthe“broad spectrum ology over the levels of causation (89 93). In portant influences on agricultural decision revolution” (BSR) (79). The BSR describes behavioral science, ultimate explanations view making, such as yield variability and labor the expansion of human diets and the in- behavior as a product of natural selection investment. These studies counter claims tensification of plant and animal exploita- (cultural and biological) whereas proximate that applications of OFT demand idealistic tion that preceded agricultural beginnings in explanations attempt to understand how purism or seek perfection in model/data fit many parts of the world. Investigations of the individuals identify challenges, make deci- (14, 50). BSR with OFT typically use the diet breadth sions, and initiate action (this being the realm Simple, well-used models of OFT like the model (DBM) and have shown a good fitof of agency). The former have been caricatured DBM (66) outline the logic of individual, model predictions with empirical data from as evolutionary “why” questions whereas the time-sensitive decisions, and, because indi- archaeological sites in southwest Asia, the latter have been equated with questions of viduals make decisions in different contexts, Neotropics, and elsewhere (e.g., refs. 6, 62, “how.” Many see the conflation of these the model anticipates behavioral variation. and 80–87). Research shows that dietary issues as a general challenge to evolutionary The extent of this variation in time and space shifts from higher- to lower-ranked plant and theory (93) whereas others recognize that both is exactly what we find interesting, and relev- animal resources, or a more even dependence are important to understanding the structure

6174 | www.pnas.org/cgi/doi/10.1073/pnas.1308938110 Gremillion et al. Downloaded by guest on September 28, 2021 of biological, cultural, and socio-ecological explanations drawn from niche construction EE offers a systematic way to evaluate the PERSPECTIVE change (42, 92). theory (NCT) (94). NCT highlights the motivations for niche-construction activities SPECIAL FEATURE: HBE models, specifically the contingency multiple kinds of feedback that flow to the such as landscape management and its in- models such as the DBM, seek ultimate niche constructor and to other populations tensification. In fact, applications of behav- explanations by identifying optimal solutions whose environments its niche-constructing ioral ecological models in archaeology often to immediate problems. They assume that activity influences (e.g., ref. 95). In doing so, highlight the active role of humans in shap- organisms possess an evolved general capac- NCT makes it possible to better characterize ing their habitats (62, 80, 97, 108). Assertions ity to achieve these optimal solutions, at least the ecological context in which optimality to the contrary (50) neglect a substantial on average. Although it is true that models must be assessed. That hunter–gatherers did amount of literature on the subject. Critically, like the DBM rest heavily on ultimate rather (and do) modify landscapes and resources for HBE permits an evaluation of the overall than proximate explanation, this trait is no calculated gain is inarguable (96–98); how- return rates associated with resource acqui- reason to reject them. Indeed, users of the ever, the extent and scope of this practice sition, which must include the costs of model regularly acknowledge that it is silent areamatterofconsiderabledebate(49).In investing in land management as well as about proximate causes while recognizing the case of domestication and agriculture, the returns of a managed landscape. EE that many such causes exist. Both proximate landscape modification has long been recog- (including HBE and OFT) can be useful and ultimate causes shape evolution, and the nized as an important phenomenon (99– for understanding the evolution of land- study of evolution must be able to identify 101). However, we do not agree with the scape management practices and other where and when the feedbacks between contention that OFT models cannot accom- forms of niche construction and for eval- “hows” and “whys” are reciprocal (92). The modate dynamic environments and argue uating the relationship between land, la- HBE modeling format makes this step pos- that EE is well-positioned to support and bor, and resource productivity. sible by being agnostic about proximate complement NCT. We see a problem with using NCT as causes, and testing hypotheses premised on Although environmental stasis might be a “single unifying approach for integrating ultimate causation. This agnosticism does not necessary to some aspects of model building consideration of human and non-human deny human agency but notes that agency, (e.g., the contingency of the DBM assumes modification of ecosystems” (ref. 50, p. 265) practice, and habitus (for example) are ulti- a static prey ranking), the simple models of because it does not systematically address mately constrained by natural selection, at foraging theory are equally applicable to dy- how or why humans alter their resource- least over time. Because the models are not namic environments (102–104). For exam- acquisition patterns. There is little doubt that absolute covering-law statements, they can ple, the decision contingency of the DBM intensive human activity modifies ecosystems accommodate feedbacks between proximate (66) is based on the marginal rate of return in diverse ways that nonetheless follow a and ultimate causes when, for example, nat- established by two important inputs: en- limited set of pathways (50). However, a ural selection (the ultimate cause) shapes the counter rate (a function of prey abundance) more intriguing question is why altered way people make decisions (the proximate and postencounter return rate (a function of ecosystems engender or permit novel because), and those decisions further shape the prey value offset by the costs of acquiring and havioral strategies in some cases but not environment of selection. processing it). Data for the latter define the others. Indeed, if hunter–gatherer land- Whether or not the domestication of plants, ranking of different prey, which is fixed until scape modification (including intentional animals, people, or landscapes entails mor- either the size (or total nutritional value) of enhancement of resource abundance) is a phological, genetic, or associational change, by the prey changes, or the energetics of cap- common feature of hunter–gatherer adap- design (i.e., “agency”) or by accident, abruptly turing and processing it do. Thus, for each tive systems, why do so few cases generate or gradually, understanding the process on the decision contingency there might be a differ- the environment of selection that yields human side of things requires an archaeology ent prey ranking, and, if so, the “optimal” diet domestic plants and animals? Clearly, more that tracks change in human behavior. Ar- in that instance may be different from during is at work than the pervasiveness of niche chaeological data typically say little about the previous instance. The standard logic of construction: insights from NCT require the individuals but speak volumes about individ- the DBM is that lower-ranked prey enter the evolutionary logic of human decision mak- uals in aggregate. HBE provides the theoretical diet as the abundance of higher-ranked prey ing in tandem with a tighter consideration foundation for understanding aggregate pat- declines; increasing abundance of lower- of the interactions between multiple taxa in terns as the product of individual decisions. ranked prey has only a minimal effect on the novel settings to explain the origins of ag- Indeed, archaeological data are best viewed as decision to include them in the diet (105, riculture. Without a theory of organismal fi the products of traits most common across 106). However, environmental modi cation behavior, such as HBE, the alternatives imply a population of individuals; change in the may affect more than just absolute abun- either (i) that humans invented agriculture spatial and temporal patterns of archaeolog- dance and rates of encounter: change in the and its long-term consequences knowingly ical data reflects change in the distribution or distribution of taxa also affects postencounter (as some advocates of human agency would frequency of traits within and among pop- return rates that, in turn, affect the ranking of have us believe), or (ii) that agriculture ulations, through space and time. different prey taxa. Harvesting and replanting emerged almost randomly as a function of can raise the density of plants within a patch, dynamic feedbacks between genes, environ- Ecological Models Accommodate Envi- which can alter the ranking of the resource ment, and the interactions between them ronmental Change. Some critics of OFT (by reducing the costs of collecting the seeds) (according to the strict version of NCT). suggest that it assumes an unchanging envi- and boost its dietary importance (which may ronment (refs. 14 and 50, p. 257). They argue have been constrained initially by scarcity) Integration of Evolutionary Frameworks that OFT models do not account for change (107). In other cases, selective, intensive har- Evolutionary theory as applied to archaeo- in the environment of selection that creates vests may reduce the body size of the average logical questions such as OA remains diverse, domestic plants and animals and suggest individual prey, again affecting the prey as expected for a relatively new but maturing that they should be discarded in favor of ranking and therefore the optimal decision. field. However, the various approaches share

Gremillion et al. PNAS | April 29, 2014 | vol. 111 | no. 17 | 6175 Downloaded by guest on September 28, 2021 a commitment to evolutionary explanation standing of domestication will be those that synthesis and, more recently, evolutionary even though they emphasize different pro- acknowledge that humans are subject to the and ecological developmental biology) have cesses, questions, and types of causation. EE forces of evolution. We find little justification earned the status of basic assumptions. The (inclusive of HBE and OFT) (61, 64, 109), for the argument that, of all species, only reality of evolution by natural selection is niche construction theory (19, 26), and Homo sapiens is capable of transcending such a core idea, arguably the most important – models of cultural transmission and gene theseforces.Wealsoadvocatetheuseof one for OA research. It unifies approaches – culture coevolution (19, 110 112) all have the theoretical frameworks that serve to link that use different analytic tools and methods fi potential to link empirical ndings to a well- testable hypotheses to ultimate explanations and target different kinds of questions. established body of knowledge, but they tar- for human behavior. However, we also wish The research programs that cluster under get different implications of selection. OFT to make it clear that, in doing so, we are not the umbrella of evolutionary theory have predicts how organisms might mobilize advocating any approach that ignores the generated, and continue to generate, testable evolved decision-making mechanisms to distinctiveness of cultural evolution or the hypotheses that contribute to our un- maximize utility in the face of conflicting great behavioral plasticity that characterizes derstanding of OA. Of these, HBE and OFT needs; niche construction theory asks how human interactions with the natural world. environmental engineering sets up pathways Nor are we in favor of glossing over historical have a longstanding record of productivity, of ecological inheritance that affect the fitness particulars in the interests of advancing and, more recently, models of cultural in- of the organism and its descendants; cultural general theories of change. heritance have begun to show considerable transmission theory asks how patterns of Calls to purge OA research of HBE or promise. Although there is no way of pre- social learning peculiar to our species gener- OFT fail to distinguish between levels of dicting which evolutionary perspectives will ate patterns of descent and divergence in theory and their respective epistemological prove most useful in the long run, one thing cultural lineages. roles. In OA research, certain core elements is certain: a retreat from theory in favor of Debates over the relative utility of these of Darwinian evolutionary theory (as ad- particularism will ensure that none of them approaches have been helpful in distinguish- vanced and amended by the neo-Darwinian achieve their full potential. ing between fundamental incommensur- abilities and superficial differences, thereby moving us closer to synthesis. We agree with 1 Denham TP, et al. (2003) Origins of agriculture at Kuk Swamp in 18 Mesoudi A, Whiten A, Laland KN (2006) Towards a unified “ the highlands of New Guinea. Science 301(5630):189–193. science of cultural evolution. Behav Brain Sci 29(4):329–347, Bettinger and Richerson that Darwinian 2 Denham T (2011) Early agriculture and plant domestication in New discussion 347–383. theory accommodates a virtually limitless Guinea and island Southeast Asia. Curr Anthropol 52(Suppl 4): 19 O’Brien MJ, Laland KN (2012) Genes, culture, and agriculture: An range of interesting, fruitful projects. It is S379–S395. example of human niche construction. Curr Anthropol 53:434–470. 3 Piperno DR, Ranere AJ, Holst I, Iriarte J, Dickau R (2009) Starch 20 Richerson PJ, Boyd R, Henrich J (2010) Colloquium paper: Gene- perhaps this diversity that causes us some- grain and phytolith evidence for early ninth millennium B.P. maize culture coevolution in the age of genomics. Proc Natl Acad Sci USA times to forget the basic commonalities that from the Central Balsas River Valley, Mexico. Proc Natl Acad Sci USA 107(Suppl 2):8985–8992. 21 unite the Darwinian enterprise”(ref. 112, p. 106(13):5019–5024. Harris M (1968) The Rise of Anthropological Theory (Thomas 4 Dillehay TD, Rossen J, Andres TC, Williams DE (2007) Preceramic Crowell, New York). 22 fl 222). To declare that any of these theoretical adoption of peanut, squash, and cotton in northern Peru. Science Smith BD (2009) Core conceptual aws in human behavioral – perspectives is fatally flawed (14, 45) closes 316(5833):1890–1893. ecology. Commun Integr Biol 6:533 534. 23 thedooronpotentiallyproductivealliances 5 Iriarte J (2007) New perspectives on plant domestication and the Zeder MA, Smith BD (2009) A conversation on agricultural development of agriculture in the new world. Rethinking Agriculture: origins: Talking past each other in a crowded room. Curr Anthropol – among them. Archaeological and Ethnoarchaeological Perspectives, eds 50:681 691. 24 For example, we believe that NCT can Denham T, Iriarte J, Vrydaghs L (Left Coast Press, Walnut Creek, CA), Vrydaghs L, Denham T (2007) Rethinking agriculture: Introductory thoughts. Rethinking Agriculture: Archaeological and pp 167–188. continue to move forward by joining forces Ethnoarchaeological Perspectives, eds Denham T, Iriarte J, 6 Piperno DR (2011) The origins of plant cultivation and Vrydaghs L (Left Coast Press, Walnut Creek, CA), pp 1–15. with other evolutionary programs. Comple- domestication in the New World tropics: Patterns, process, and new 25 Zeder MA (2006) Central questions in the domestication of mentarity between NCT and EE has already developments. Curr Anthropol 52(Suppl 4):S453–S469. plants and animals. Evol Anthropol 15:105–117. 7 Iriarte J (2009) Narrowing the gap exploring the diversity of early been illustrated in case studies of New World 26 Smith BD (2007) Niche construction and the behavioral context food-production economies in the Americas. Curr Anthropol 50(5): – of plant and animal domestication. Evol Anthropol 16:188–199. hunter gatherers and agriculturalists (108). 677–680. 27 Smith BD (2001) Documenting plant domestication: The 8 Barton L, et al. (2009) Agricultural origins and the isotopic identity Broughton et al. (ref. 108, p. 371) note that consilience of biological and archaeological approaches. Proc Natl of domestication in northern China. Proc Natl Acad Sci USA 106(14): EE “focuses on how behavioral adjustments Acad Sci USA 98(4):1324–1326. 5523–5528. 28 Ketelaar T (2003) The evaluation of competing approaches to changing socio-ecological conditions cre- 9 Zhao Z (2011) New archaeobotanic data for the study of the within human evolutionary psychology. Evolutionary Psychology: origins of agriculture in China. Curr Anthropol 52(Suppl 4): ate novel selective pressures that in turn drive Alternative Approaches, eds Scher SJ, Rauscher F (Kluwer, Boston), S295–S306. other changes in morphology and behavior” pp 31–54. 10 Fuller DQ, Harvey E, Qin L (2007) Presumed domestication? 29 Zeder MA (2008) The Neolithic macro-(r)evolution: and therefore “draws on the same evolu- fi Evidence for wild rice cultivation and domestication in the fth Macroevolutionary theory and the study of culture change. – tionary logic that underlies niche construc- millennium BC of the Lower Yangtze region. Antiquity 81:316 331. J Archaeol Res 17(1):1–63. 11 tion theory.” Others (e.g., ref. 113) illustrate Fuller DQ, et al. (2009) The domestication process and 30 Zeder MA (2008) Domestication and early agriculture in the domestication rate in rice: Spikelet bases from the Lower Yangtze. Mediterranean Basin: Origins, diffusion, and impact. Proc Natl Acad how NCT-based hypotheses can explain – Science 323(5921):1607 1610. Sci USA 105(33):11597–11604. 12 technological change across the transition to Bettinger RL, Barton L, Morgan C (2010) The origins of food 31 Bar-Yosef O (2011) Climatic fluctuations and early farming in agriculture. O’Brien and Laland (19) make production in North China: A different kind of agricultural revolution. West and East Asia. Curr Anthropol 52(Suppl 4):S175–S193. – astrongcasethatgene–culture coevolution Evol Anthropol 19(1):9 21. 32 Belfer-Cohen A, Goring-Morris AN (2011) Becoming farmers: 13 Bettinger RL, et al. (2010) The transition to agriculture at The inside story. Curr Anthropol 52(Suppl 4):S209–S220. theory and NCT strengthen and support Dadiwan, People’s Republic of China. Curr Anthropol 51(5):703–714. 33 Belfer-Cohen A, Goring-Morris AN (2009) For the first time. Curr explanations of adaptive complexes related to 14 Zeder MA (2012) The Broad Spectrum Revolution at 40: Anthropol 50:669–671. Resource diversity, intensification, and an alternative to optimal 34 Flinn M (1997) Culture and the evolution of social learning. Evol agriculture, such as adult lactose tolerance foraging explanations. J Anthropol Archaeol 31:241–264. Hum Behav 18:23–67. and persistence of sickle-cell disease. 15 Hegmon M (2003) Setting theoretical egos aside: Issues and 35 Henrich J, McElreath R (2003) The evolution of cultural evolution. theory in North American archaeology. Am Antiq 68:213–243. Evol Anthropol 12:123–135. Conclusions 16 Ellen R (2010) Theories in anthropology and “anthropological 36 Nettle D (2009) Beyond nature versus culture: Cultural variation fi theory”. J R Anthropol Inst 16:387–404. as an evolved characteristic. J R Anthropol Inst 15:223–240. We believe strongly that the most signi cant 17 Boyd R, Richerson PJ (1985) Culture and the Evolutionary 37 Jones BD (1999) Bounded rationality. Annu Rev Polit Sci archaeological contributions to our under- Process (Univ of Chicago Press, Chicago). 2:297–321.

6176 | www.pnas.org/cgi/doi/10.1073/pnas.1308938110 Gremillion et al. Downloaded by guest on September 28, 2021 38 Simon HA (1947) Administrative Behavior (Macmillan, New 66 MacArthur RH, Pianka ER (1966) On optimal use of a patchy 89 Mayr E (1961) Cause and effect in biology. Science 134(3489): PERSPECTIVE York). environment. Am Nat 100:603–609. 1501–1506. SPECIAL FEATURE: 39 Hodder I (1990) The Domestication of Europe: Structure and 67 Smith BD (2006) Human behavioral ecology and the transition to 90 Tinbergen N (1963) On aims and methods of ethology. Contingency in Neolithic Societies (B. Blackwell, Oxford). food production. Behavioral Ecology and the Transition to Z Tierpsychol 20:410–433. 40 Cauvin J (2000) The Birth of the Gods and the Origins of Agriculture, eds Kennett DJ, Winterhalder B (Univ of California Press, 91 West-Eberhard MJ (2003) Developmental Plasticity and Agriculture, trans Watkins T (Cambridge Univ Press, Cambridge, UK). Berkeley, CA), pp 289–303. Evolution (Oxford Univ Press, Oxford). 41 Gardner A (2011) Action and structure in interpretive 68 Levins R (1966) The strategy of model-building in population 92 Laland KN, Sterelny K, Odling-Smee J, Hoppitt W, Uller T (2011) archaeologies. Evolutionary and Interpretive Archaeologies: A biology. Am Sci 54:421–431. Cause and effect in biology revisited: Is Mayr’s proximate-ultimate Dialogue, eds Cochrane EE, Gardner A (Left Coast Press, Walnut 69 Richerson PJ, Boyd R (1987) Simple models of complex dichotomy still useful? Science 334(6062):1512–1516. Creek, CA), pp 63–82. phenomena: The case of cultural evolution. The Latest on the Best: 93 Scott-Phillips TC, Dickins TE, West SA (2011) Evolutionary theory 42 Smith EA (2013) Agency and adaptation. Annu Rev Anthropol Essays on Evolution and Optimality, ed Dupre J (MIT Press, and the ultimate-proximate distinction in the human behavioral 42:103–120. Cambridge, MA), pp 27–52. sciences. Perspect Psychol Sci 6:38–47. 43 Ingold T (2000) The Perception of the Environment: Essays in 70 Winterhalder B (2002) Models. Darwin and Archaeology: A 94 Odling-Smee J, Laland KN, Feldman W (2003) Niche Livelihood, Dwelling and Skill (Routledge, London). Handbook of Key Concepts, eds Hart JP, Terrell JE (Bergin and Garvey, Construction (Princeton Univ Press, Princeton). 44 Joseph S (2000) Anthropological evolutionary ecology: A Westport, CT), pp 201–224. 95 Odling-Smee J, Laland KN (2011) Ecological inheritance and critique. J Ecol Anthropol 4:6–30. 71 Catania KC (2012) Evolution of brains and behavior for cultural inheritance: What are they and how do they differ? Biol 45 Cochrane EE, Gardner A, eds (2011) Evolutionary and optimal foraging: A tale of two predators. Proc Natl Acad Sci USA Theory 6(3):220–230. Interpretive Archaeologies: A Dialogue (Left Coast Press, Walnut 109(Suppl 1):10701–10708. 96 Anderson MK (2005) Tending the Wild: Native American Creek, CA). 72 Lai Y, Chen J-H, Lee L-L (2011) Prey selection of shell-invading knowledge and the management of California’s natural resources 46 Winterhalder B, Smith EA (1992) Evolutionary ecology and the leech as predicted by optimal foraging theory with consumption (Univ of California Press, Berkeley, CA). social sciences. Evolutionary Ecology and Human Behavior, eds success incorporated into estimation of prey profitability. Funct Ecol 97 Bliege Bird R, Bird DW, Codding BF, Parker CH, Jones JH (2008) Smith EA, Winterhalder B (Aldine de Gruyter, New York), pp 3–24. 25:147–157. The “fire stick farming” hypothesis: Australian Aboriginal foraging 47 Nitecki MH, Nitecki DV (1992) History and Evolution (State Univ 73 Danial-Valcroze T, Lesage V, Giard J, Michaud R (2011) Optimal strategies, biodiversity, and anthropogenic fire mosaics. Proc Natl of New York Press, Albany, NY). foraging theory predicts diving and feeding strategies of the largest Acad Sci USA 105(39):14796–14801. 48 Richerson PJ, Boyd R (1997) Built for speed, not for comfort: marine predator. Behav Ecol 22:880–888. 98 Smith BD (2007) Behavior: The ultimate ecosystem engineers. Darwinian theory and human culture. Philosophica 60:13–43. 74 Sayers K, Norconk MA, Conklin-Brittain NL (2010) Optimal Science 315(5820):1797–1798. 49 Lightfoot KG, Cuthrell RQ, Striplen CJ, Hylkema MG (2013) foraging on the roof of the world: Himalayan langurs and the 99 Anderson E (1952) Plants, Man, and Life (Univ of California Rethinking the study of landscape management practices among classical prey model. Am J Phys Anthropol 141(3):337–357. Press, Berkeley, CA). hunter-gatherers in North America. Am Antiq 78(2):285–301. 75 Levi T, Lu F, Tu DS, Mangel M (2011) The behaviour and diet 100 Anderson E (1956) Man as a maker of new plants and new 50 Smith BD (2011) A cultural niche construction theory of initial breadth of central-place foragers: An application ot human hunters plant communities. Man’s Role in Changing the Face of the Earth,ed domestication. Biol Theory 6:260–271. and Neotropical game management. Evol Ecol Res 13:171–185. Thomas WL (Univ of Chicago Press, Chicago), pp 763–777. 51 Gray RD (1987) Faith and foraging: A critique of the “paradigm 76 Lupo KD, Fancher JM, Schmitt DN (2012) The taphonomy of 101 Lewis HT (1972) The role of fire in the domestication of plants argument from design”. Foraging Behavior, eds Kamil AC, Krebs JR, resource intensification: Zooarchaeological implications of resource and animals in Southwest Asia: A hypothesis. Man (Lond) 7(2): Pulliam HR (Plenum, New York), pp 69–140. scarcity among Bofi and Aka forest foragers. J Archaeol Method 195–222. 52 Pierce GJ, Ollason JG (1987) Eight reasons why optimal foraging Theory 20:420–447. 102 Belovsky GE (1988) An optimal foraging-based model of theory is a complete waste of time. Oikos 49:111–118. 77 Dwairy M, Dowell AC, Stahl J-C (2011) The application of hunter-gatherer population dynamics. J Anthropol Archaeol 7(4): 53 Boyer P (1995) Ceteris paribus (all else being equal). How Things foraging theory to the information searching behaviour of general 329–372. Are: A Science Tool-Kit for the Mind, eds Brockman J, Matson K practitioners. BMC Fam Pract 12:90–98. 103 Winterhalder B (1986) Diet choice, risk, and food sharing in (William Morrow, New York), pp 169–175. 78 Robinson ME, McKillop HI (2013) Ancient Maya wood selection a stochastic environment. J Anthropol Archaeol 5:369–392. 54 Yengoyan AA (2004) Anthropological History and the Study of and forest exploitation: A view from the Paynes Creek salt works, 104 Winterhalder B, et al. (1988) The populations ecology of Hunters and Gatherers: Cultural and Non-Cultural Hunter-Gatherers Belize. J Archaeol Sci 10:3584–3595. Hunter-Gatherers and their prey. J Anthropol Archaeol 7:289–328. in History, Archaeology and Anthropology, ed Barnard A , Berg, New 79 Flannery KV (1969) Origins and ecological effects of early 105 Pyke GH, Pulliam HR, Charnov EL (1977) Optimal foraging: A York), pp 57–66. domestication in Iran and the Near East. The Domestication and selective review of theory and tests. Q Rev Biol 52:137–154. 55 Henrich J (2002) Decision making, cultural transmission, and Exploitation of Plants and Animals, eds Ucko PJ, Dimbleby GW 106 Winterhalder B (1981) Optimal foraging strategies and hunter- adaptation in economic anthropology. Theory in Economic (Aldine, Chicago), pp 73–100. gatherer research in anthropology: Theory and methods. Hunter- Anthropology, ed Ensminger J (Alta Mira, Walnut Creek, CA), pp 80 Piperno DR (2006) The origins of plant cultivation and Gatherer Foraging Strategies: Ethnographic and Archeological 251–295. domestication in the Neotropics: A behavioral ecological perspective. Analyses, eds Winterhalder B, Smith EA (Univ of Chicago Press, 56 Henrich J (2004) Demography and cultural evolution: How Behavioral Ecology and the Transition to Agriculture, eds Kennett D, Chicago), pp 13–35. adaptive cultural processes can produce maladaptive losses—the Winterhalder B (Univ of California Press, Berkeley, CA), pp 137–166. 107 Winterhalder B, Goland C (1997) An evolutionary ecology Tasmanian case. Am Antiq 69(2):197–214. 81 Stiner MC, Munro ND, Surovell TA (2000) The tortoise and the perspective on diet choice, risk, and plant domestication. People, 57 McElreath R (2004) Social learning and the maintenance of hare: Small game use, the broad spectrum revolution, and Paleolithic Plants, and Landscapes: Studies in Paleoethnobotany,ed cultural variation: An evolutionary model and data from East Africa. demography. Curr Anthropol 41(1):39–79. Gremillion KJ (Univ of Alabama Press, Tuscaloosa, AL), pp 123–160. Am Anthropol 106(2):308–321. 82 Stiner MC, Munro DN (2002) Approaches to prehistoric diet 108 Broughton JM, Cannon MD, Bartelink EJ (2010) Evolutionary 58 Richerson PJ, Boyd R (2005) Not by Genes Alone: How Culture breadth, demography, and prey ranking systems in time and space. ecology, resource depression, and niche construction theory: Transformed Human Evolution (Univ of Chicago Press, Chicago). J Archaeol Method Theory 9:181–214. Applications to central California hunter-gatherers and Mimbres- 59 Bird D, O’Connell JF (2006) Behavioral ecology and archaeology. 83 Jones EL (2009) Climate change, patch choice, and Mogollon agriculturalists. J Archaeol Method Theory 17:371–421. J Archaeol Res 14:143–188. intensification at Pont d’Ambon (Dordogne, France) during the 109 Bettinger RL (2006) Agriculture, archaeology, and human 60 Gremillion KJ (2004) Seed processing and the origins of food Younger Dryas. Quat Res 72:371–376. behavioral ecology. Behavioral Ecology and the Transition to production in eastern North America. Am Antiq 69:215–234. 84 Munro ND (2009) Epipaleolithic subsistence intensification in the Agriculture, eds Kennett DJ, Winterhalder B (Univ of California Press, 61 Gremillion KJ (1998) Changing roles of wild and cultivated plant southern Levant: The faunal evidence. Integrating Approaches to the Berkeley, CA), pp 304–322. resources among early farmers of Eastern Kentucky. Southeastern Study of Paleolithic Subsistence, eds Hublin J-J, Richards MJ (Springer, 110 Richerson PJ, Boyd R, Bettinger RL (2001) Was agriculture Archaeol 17:140–157. The Hague), pp 141–155. impossible during the Pleistocene but mandatory during the 62 Piperno DR, Pearsall DM (1998) The Origins of Agriculture in the 85 Munro ND (2009) Integrating inter- and intra-site analyses of Holocene? A climate change hypothesis. Am Antiq 66:387–412. Lowland Neotropics (Academic, San Diego). Epipaleolithic faunal assemblages from Israel. Before Farming: 111 Coward F, Shennan S, Colledge S, Conolly J, Collard M (2008) 63 Barlow KR (2006) A formal model for predicting agriculture Archaeol Anthropol Hunter-Gatherers. 2009(1):4. The spread of Neolithic plant economies from the Near East to among the Fremont. Behavioral Ecology and the Transition to 86 Stutz AJ, Munro ND, Bar-Oz G (2009) Increasing the resolution of northwest Europe: A phylogenetic analysis. J Archaeol Sci 35:42–56. Agriculture, eds Kennett DJ, Winterhalder B (Univ of California Press, the Broad Spectrum Revolution in the Southern Levantine 112 Bettinger RL, Richerson PJ (1996) The state of evolutionary Berkeley, CA), pp 87–102. Epipaleolithic (19-12 ka). J Hum Evol 56(3):294–306. archaeology: Evolutionary correctness, or the search for common 64 Marston JM (2011) Archaeological markers of agricultural risk 87 Starkovich BM, Stiner MC (2010) Upper Palaeolithic animal ground. Darwinian Archaeologies, ed Maschner HDG (Plenum, New management. J Anthropol Archaeol 30:190–205. exploitation at Klissoura Cave 1 in southern Greece: Dietary trends York), pp 221–231. 65 Foster HT II (2003) Dynamic optimization of horticulture among and mammal taphonomy. Eurasian Prehistory 7:107–132. 113 Collard M, Buchanan B, Ruttle A, O’Brien MJ (2011) Niche the Muscogee Creek Indians of the southeastern United States. 88 Ugan A, Simms S (2012) On prey mobility, prey rank, and construction and the toolkits of hunter-gatherers and food J Anthropol Archaeol 22:411–424. foraging goals. Am Antiq 77(1):179–185. producers. Biol Theory 6:251–259.

Gremillion et al. PNAS | April 29, 2014 | vol. 111 | no. 17 | 6177 Downloaded by guest on September 28, 2021