5

Cooperation, Conflict, and Niche
Construction in the Genus Homo

Agustín Fuentes

Introduction
While it is important to acknowledge the power of a focus on selection and extant
Neo-Darwinian theory, we should also emphasize that there is more to evolution than
that. Utilizing the traditional concept, a trait-based selection scenario, may be too limiting
when thinking about, or modeling, the evolution of complex social systems/organisms.
Evolution is ongoing and the contexts, patterns, and mechanisms of evolutionary processes
are what should interest us most. This is an argument against the common particularly
static take on selection and evolution . . . it is not so much that things have evolved, but
that they are evolving.
We are in the midst of significant enhancements in complexity and diversity in evolu-
tionary theory, with the role for behavioral modification of social and ecological spaces, and
their inheritance, becoming a key factor. Our grasp of patterns and contexts of selection
and the ways in which social, epigenetic and developmental interactors affect outcomes is
growing by leaps and bounds. We also are increasingly realizing that social and experiential
contexts shape bodies and behavior, affecting trajectories in more substantial manners than
previously envisioned. In this chapter I will review current proposals for integrating per-
spectives from niche construction and multi-inheritance into investigations, models, and
explanations for the evolution of human behavior.
Mary Jane West-Eberhard’s broad overview (2003) of developmental plasticity and
evolution led her to suggest that plasticity is one of the key factors for our understand-
ing of adaptive evolution. She argues, like many other evolutionary biologists, that reduc-
ing the processes of development and evolutionary change to genomic levels is not always

and con- vey more than material descriptions. & Relyea. life history. physiology. Padilla. or even marked. Jablonka and Lamb argue for adding a per- spective wherein three other inheritance systems can also have causal roles in evolutionary change. This allows for the acquisition and reproduction of a variety of behaviors. Their main point being that practitioners of traditional Neo-Darwinian approaches focus on one system of inheritance: the genetic system of inheritance. 2005). for example). contain a high density of information. such models may be better attuned to the actual interactions of systems. the favoring of close relatives due to their high degree of shared genotype. This moves past standard Neo-Darwinian approaches. 2005). morphology. Morgan. thus acquisition of evolutionarily relevant behavioral patterns can occur through socially mediated learning. in the sense that which variants are inher- ited and what final form they assume depend on various filtering and editing processes that occur before and during transmission ( Jablonka & Lamb. This transmission of information occurs without having any linkage to genetic systems that natural selection (in a Neo-Darwinian view) can target. this perspective forces an evolutionary concern with the way in which human bod- ies and behavioral and symbolic systems construct and interact with social and ecological . Recent reviews define basic phenotypic plasticity as “the production of multiple phe- notypes from a single genotype. Research into modeling this plasticity. C o o p er at i o n . These other systems are the epigenetic. Epigenetic inheritance is found in all organisms. a n d Ni c h e C o n st ru c t i o n 79 possible or preferable. However. Specifically in terms of human evolu- tion. perceptions. but adaptive. and symbolic inheritance is found only in humans. its potential adaptive value and contexts. This phenotypic plasticity and its relation to ecologies and evolutionary patterns is of core interest in evolutionary theory. Information is transferred from one generation to the next by many interacting inheritance systems. Sutlan. Variation is also constructed. Many organisms transmit information via behavior. 2005). and symbolic inheritance sys- tems. depending on environmental conditions” (Miner. a “new” new synthesis in how we model evolution. Biologists Eva Jablonka and Marion Lamb (2005) call for a renovation in evolution- ary theory. C o n fli c t. genetic change. However. and demography and that this plastic- ity can occur in both individually and inter-generational contexts ( Jablonka and Lamb. Models using this system become more complex than the general reductionist models of Neo-Darwinian behavioral theory (such as kin selection. These analyses demonstrate that evolved plasticity in development enables the evolution of new or variant. and its ecological impact all suggest that phenotypic plasticity is a significant factor for many organisms’ evolutionary histories and current behavior/morphology. Symbolic inheritance comes with language and the ability to engage in information transfer that can be temporally and spatially complex. more important than the basic definition is the evidence that a range of organisms express phenotypic plasticity via changes in behav- ior. behavioral. phenotypes without substantial. They argue for recogni- tion of “evolution in four dimensions” rather than a focus on just one. growth. and beliefs that are potentially beneficial for individual humans and populations and that have no genetic basis or linkage. behavioral inheritance in most.

Niche construction creates feedback within the evolutionary dynamic. 2007). contributes to changes over time in the dynamic relationship between organisms and environments. Niche construction is the building and destroying of niches by organisms and the synergistic interactions between organisms and environments. but the transmission of this knowledge it itself dependent on preexisting information acquired through genetic evolution. Fuentes et. Niche con- struction reflects a synthesis of ecological. Odling-Smee et al.g. Tehrani & Odling-Smee. Niche construction impacts/alters energy flows in ecosystems through ecosystem engineer- ing creating an ecological inheritance and. and niche construction in general can occur via cultural means. and Feldman (2003) proposed niche construction as a significant evolutionary force. contexts. and taking from the extended phenotype concept of Richard Dawkins (1982). Brown. Odling-Smee. They state that humans are the “ultimate niche constructors” and that adding niche construction to attempts to understand human systems makes such attempts more complicated (bypass- ing more simplistic Neo-Darwinian adaptationist accounts).g. each of which can impact patterns. Kendall et al. these systems interact with epigenetic and genetic systems.. 2010. 2005. Odling-Smee et al. (2003) explicitly state that ecological inheritance. Day. 2011). and social niches rather than treating them as discreet spheres (e. like natural selection. (2003) propose a specific model for human genetic and cultural evolution that they call a Tri-inheritance vision model. Laland & Odling-Smee. Bolhuis. Gamble & Gowlett. Richerson & Boyd. complex ontogenetic processes. human behavior results from information acquiring processes at three levels: population genetic processes. 2003. Building on the work of Richard Lewontin (1983) and earlier perspectives of Ernst Mayr (1963) and Conrad Waddington (1959). via material culture. and on unrelated populations sharing the same landscape. Wyczalkowski. They state “Much of human niche construction is guided by socially learned knowledge and cultural inheritance.. biological. Laland. & Laland. then the role of social/symbolic and ecological inheritance and intra.. ontogenetic processes and cultural processes. on their descendants. and structure of natural selection.. 2010. 2011.. & MacKinnon. They see cultural processes as providing a particularly robust vehicle for niche construction. as shown in Table 5. 2011.1. pp. Wells & Stock.and inter- group interactions and relationships and their impacts on ecosystems come to the forefront in our examinations of evolutionary trajectories and processes. or prior social learning” (Odling-Smee et al. 2010: Kendall. Richardson. Henrich. If we see niche construction and multiple lines of inheritance as a core to evolutionary process. such that organisms engaged in niche construction modify the evolutionary pressures acting on them. . In the model. This perspective blurs any clear prioritization in inheritance systems and forces a move away from approaches that are limited to either social or biological foci. Dunbar. al.80 Eco l o gi c a l a n d Evo lu t i o na ry Mo d el s niches and how. in turn. This is borne out by a diverse array of recent work (e. 260–261). Niche construction in humans emerges from all three of these processes. 2011. 2003: Fuentes.

Even when moving into new areas. and addition to the developmental cultural processes biological and ecological factors throughout the course of life history Basic premise • there is more to Evolution is not a matter of • ecosystem engineering heredity than genes organisms or populations being • organisms modify their. and between genic. predation risk. including • some acquired time. . epigenetic. development contributes to changes over time instruction as well as as construction.1 Summaries of Main Points of Emerging Perspectives in Evolutionary Theory Evolution in four Developmental systems theory Niche construction dimensions Focus of selection Genetic. and • some hereditary molded by their environments other. for information is determination by multiple subsequent populations inherited causes. Understanding human evolution requires assessing the interactive and mutually mutable relationship humans have with their social and struc- tural ecologies. 2009). cultural information acquiring variations. Constant constructing—and Tri-inheritance vision model causes for epigenetic. distributed in the dynamic relationship selection control. We must accept the possibility that selection pressures can be modified as they are occurring and that human response to selective challenges will always fit the expected parameters of the selective force. context sensitivity • is a process. C o o p er at i o n . and the extraction of nutrition from complex food sources had impacts on the devel- opment of human cognition and adaptive strategies in a diverse array of areas (Gowlett. and ecology interaction (niche)—a symbolic systems behavioral factors focus on Phenogenotypes Main underlying Combination of genic. and intergroup contexts in ontogenetic processes. behavioral being constructed (TIV) wherein human evolution of and symbolic by—demography. extended to natural selection. C o n fli c t. social behavior results from human behavior inheritance systems interactions. in addition • evolutionary change and contingency. Wrangham. population genetic processes. 2010. and manipulation processes at three levels: of the environment in intra. Outcome of complex interactions Individual and individual-local behavioral. organisms’ selective variations are but of organism-environment environments nonrandom in origin systems changing over • ecological inheritance. For example. This involves: joint modified selection pressures. and evolution as between organisms and construction: environments (niches) Niche Construction and Human Evolution The ability of humans to modify their social and physical surroundings is central to any explanation of human behavior. epigenetic. behavioral innovation via the use of controlled fire as a response to pressures exerted by climate stress. That is. Humans generally exist in a place where there have been humans previously. human toolkits (our bodies and minds) might result in innovation that adds elements into a system that standard evolutionary approaches cannot foresee or do not normally include. The social and ecological landscapes are impacted by the previous generations so that subse- quent human generations inherit a known and human-altered landscape and ecology and a substantial amount of information about living in that place (which increases with the advent of even rudimentary language and tool use). a n d Ni c h e C o n st ru c t i o n 81 TABLE 5. that can result from inheritance.

aspects critical to niche construction. particularly the evolution of the genus Homo from the Pleistocene on we need to focus both on continuities and discontinuities in our evolutionary history. Gamble & Gowlett. humans can develop multiple responses to the same challenges and potentially share them across individuals and possibly even groups. This provides humans with more ways to respond to challenges. most hypotheses for the evolution of human behavior relied strictly on natural selection as the only significant evolutionary force in the structuring of human behavior. than other organisms.82 Eco l o gi c a l a n d Evo lu t i o na ry Mo d el s humans carry social and material knowledge of their group with them and pass it on to the generations in the new location. Mackinnon & Fuentes. & Gray. 2005) focuses on the possibility that selection acts on human cultural behavior.” monumental shifts in selection events/ responses that catapult a lineage from one stage to the next. Is it possible that in the mainstream approaches to examining human evolution we have overlooked some core aspects that might be right under our noses. Dual inheritance theory (e. Fuentes & White 2012). Mainstream approaches in the inquiry into human evolution frequently look for “big moments. 1998. Until recently. Assessing human evolution really becomes a quest to figure out why Homo succeeded when all the other hominins went extinct and the hominin’s sibling genera (African and Asian great apes) never really made the leap to a broadly successful suite of adaptations (Hare. 2001).. Successful human responses can reflect a pattern of plasticity and flexibility resulting from a cohort of selection pres- sures as opposed to specific selection for a particular adaptation in response to a single selective pressure. Selection need not always be invoked to explain the innovation and spread of behavior. and with a more diverse array of means.. In understanding and investigating human evolution. 2010. Malone. phenotypic. Given this pat- tern of shared behavioral plasticity humans may be able to respond at quicker rates than most complex organisms when faced with strong selective challenges. we should also include possible impacts from gene flow and drift in genetic. It is most likely that the majority of human responses that result in behavioral change over time are not optimal. Small and/or complex multifaceted changes over time may not leave such dramatic signals and thus a focus on major morphological shifts or the model- ing of massive single event behavioral shifts might not be the best way to approach our evolution. and it is highly likely that patterns of flow and drift can impact behavior as well as geno- typic factors. This suggests that explanations that focus on the link of a particular behavior to a specific adaptive outcome may be poor models for human behavioral evolu- tionary processes. When it is invoked we should also be prepared to include models that accept selection as acting on levels beyond a selfish gene or selfish individual focus (as highlighted by Dunbar. behavioral and even symbolic inheritance? . Odling-Smee et al. Oyama. Richerson & Boyd. 2011. However. even if they do result in adaptation.g. Griffiths. 2011. and behavioral responses. Sober & Wilson. This might not always be the best path to take. 2003.

Sussman and Cloninger (2011). social networks. in turn. and possibly inter-. such as the explanations proposed by Lovejoy (2009) for pair-bonding and monogamy in early hominins.) than mod- eling dyadic interactions between group members (Dunbar. For example. Gamble & Gowlett. Do evolutionary processes produce the best outcomes or ones that are just good enough? Largely. This does not mean that specific traits and sexual selection are not at play and important. argues for male carrying as a way to reduce reproductive costs on females and increase infant survivorship and suggest that this is part of a network of adaptations that characterize caretaking of high-cost infants in members of the genus Homo. the latter. especially in the areas of aggression and cooperation. but simply that more comprehensive system processes might also be central to our evolutionary patterns. forcing a focus on more than dyadic interactions as the basis of their arguments. Gamble. First. and others. 2010. but also might lead to enhanced explanatory models and insight into modern human behavior. C o n fli c t. there is emerging consensus that approaches to reconstructing behavior by Pleistocene members of the genus Homo are bet- ter served by looking at group sizes. Direct and continuous selection pressure . This implies a shift away from specific trait and sexual selection models as the primary explanatory modes in the evolution of human behav- ior. respectively. from an evolutionary sense. Before moving on to present a model for the patterns I suggest. C o o p er at i o n . I argue that we have ignored some important possibilities that emerge from thinking about behavior within and between groups that might provide not only additional information about human evolutionary trends. we need to acknowl- edge a few potential theoretical and ideological stumbling blocks that might affect our abilities to think about human (and primate) evolution. More specifically. Nowak and Highfield (2011). group behavior might hold some important answers/concepts to better model human evolution. etc. use of fire. and intergroup interactions and their relationships to changes in the archeological record (tool types. I propose that a more comprehensive and inclusive look at intra-. a n d Ni c h e C o n st ru c t i o n 83 Cooperation and Reciprocity as Core to Human Success Following Dunbar. The first is the over-emphasis on the concept of optimality. Both of these examples use specific behavioral traits in traditional fitness assessments. Expanding outside the “pair focus” might enable us to get a better grasp on why we do what we do. in as core factors in human evolution. the male-female unit: we do not live in dyads nor have dyads been the main unit of interaction for humans for millions of years (although pair bonds are important for humans). Fuentes 2009). but they rest the efficacy of the model in the context of multi-individual social networks. Hrdy interweaves specific trait-based assumptions about reproductive success and connects their enhancement to a specific scenario of multi-individual allocare. and Gowlett (2010). we have to move away from the notion that the basal unit of analysis for human behavior and interaction is the pair. This is illustrated by Hrdy (2005) and Gettler (2010) when they argue for cooperative breeding and extended male participation in child-care. Gettler.

Sussman & Cloninger. There is mounting evidence from the fossil. If we are always assuming that systems move toward an optimum and that over evolutionary time. but none of these traits occur in a vacuum and it behooves us to remember that optimality modeling is really just a heuris- tic tool. but might be very well suited to an organism that relies on a diverse tool kit for engaging with social and ecological landscapes. and a deleterious approach at worst when investigating human evo- lution (e. while others may not be. we might ignore the possibility that there are multiple effective outcomes for a given trait in a given system. 2011). and meerkats. we are better positioned to think in terms of multiple systems simultaneously impacting the human evolutionary trajectory. or stabilizing selection as the only relevant forces on them. and neurobiolo- gies of the primates (Bekoff 2007. e. if we are not concerned with optimality but rather with system function in the sense of evolutionary sufficiency.. complex social cooperation and reciprocity form the central modes of interac- tion in many mammalian groups (coyotes. there has traditionally been sub- stantial emphasis placed on direct competition and aggression between dyads and between groups (Fuentes.84 Eco l o gi c a l a n d Evo lu t i o na ry Mo d el s toward optimality can be modeled for any trait in question. de Waal & Brosnan. wolves. This is not unique to human evolution. For example. independent of their role and position in a given system. disruptive. Nowak & Highfield. espe- cially with the great apes. Hart & Sussman. The second way to enhance our evolutionary models and insights is to query: what is adaptive? If we are open to the possibility that a myriad of traits (both physical and behav- ioral) may not be strictly adaptive. Rilling. In this light it is important to point out that conflict/competition are not the only drivers in evolutionary systems. Potential for emergent properties and their relation to evolutionary change is important for us to consider. Not all relevant elements in the evolu- tion of systems operate with either directional. we must be wary of the benefits and drawbacks of comparative models. evolutionary histories. 2011. comparative primatological. and modeling literature that this is an incom- plete approach at best. In the investigation of human behavioral evolution. just to name a few) and are especially important in the daily lives. This can go hand in hand with a recognition of plasticity in behavioral patterns as a form of preadaptation which is not optimal. We might also make the mistake of describing traits as if they exist by themselves and can be assessed as such.. Finally. 2011. but yet still influence the functioning and potential of a given system. not a reflection of real systems’ operations. Chimpanzee analogies in human evolutions are only partial at .g. Weiss & Buchanan. 2006). we are open to more possibilities for explanation of patterns without denying the possibility that some traits may be approaching optimality in function and/or structure. We have to consider that there may be a mutual mutability in cooperative and competitive relationships and a degree of plasticity and hybridity in function in addition to our general views of adaptation and strategies. archeological. since cooperation as a core driver in the evolution of organismal and social systems can be found throughout the animal kingdom (and beyond.g. Sussman & Cloninger. 2009. 2009). 2008). whales. 2011. traits that persevere are opti- mal. However.

and humans) are highly derived and that the recent common ancestor (RCA) was largely unlike the present forms. ecological. or idealized. a n d Ni c h e C o n st ru c t i o n 85 best. C o o p er at i o n . not only those in common with other primates as has regularly been done with apes (e. Considering the available fossil. or bonobos. as the genus Homo and the genus Pan (consisting of the two species P. morphologically. we need to consider what human groups did in the past and what we continue to do. should be examined from a systems perspective. for example. the association of males (and not females) with the manu- facture and use of tools and other material technical skills including creation of material art.. 2011. are not supported by any fossil or archeological evidence before the last 10 to 20. division of labor as a baseline for the construction of evolutionary “roles” of males and females regarding conflict and cooperation. 2008). Olga & Jake. C o n fli c t. paniscus. or ecologically) for the RCA (Ferguson. Hart & Sussman. and assumed to be present in past human groups. This means that behavioral analogies need to be utilized with extreme care. Dunbar. and physiological arenas. social plasticity. 2011. and the expectation of males taking lead roles in group coordination are all assump- tions. The fossil and material record do not provide evidence for the vast majority of the modern gender patterns until very recently (Adovasio. based on modern gender roles and stereotypes. physi- ological. not to use a priori modern gender roles and the optimal. This is very noticeable when it comes to issues of aggression and the Homo-Pan comparisons. troglodytes. and P. The expectation of mini- mal male caretaking of children. There is a pattern wherein the archeological evidence and the comparative .g. and niche constructivist tendencies. Systems Approaches Human behavior. Simply put. muscle density. but that behaviorally they are quite divergent in most areas. that are frequently imposed on the human past. 2010. Gamble & Gowlett. 2011). gorillas. While we can argue for behavioral differences based on size. and aspects of repro- ductive physiology. MacKinnon & Fuentes. or chimpanzees) are extremely different in a wide array of behavioral.000 years. but in explaining the radical suc- cess of genus Homo relative to the other Hominoids we must focus on human derived traits. looking at social systems on the whole rather than as a series of dyadic con- tests or specific concordances (or lack thereof ) with optimal predictions may best explain many of the overall behavioral patterns in human and human evolution. We do need to be cautious with the comparative primatological approach as it can misdirect research foci away from important hominin features such as our extreme offspring costs. We should be careful. Hare. and the evolution of that behavior. 2007). It should be quite clear at this point that humans and the two Pan species may share some similarities. That is. we must stop envision- ing chimpanzees and bonobos as a model (behaviorally. Chimpanzees can provide some good phylogenetic comparisons (as can other primates). and morphological evidence it is highly likely that each of the African homi- noid lineages (chimpanzees/bonobos. many of the gender role assignments (from tool-making to sexuality to childcare to group leadership) found in societies today.

mainland Asia. provisioning. may have emerged as a core process in human evolution that is not best modeled as dyadic interactions or contests. or system.86 Eco l o gi c a l a n d Evo lu t i o na ry Mo d el s primatological approach do not definitively point to one specific way of being successfully human. 2003) or maternally related males (i. rather it reflects the fact that human soci- eties are based on extensive and extremely complex systems of cooperation and mutual interreliance on one another. (2004) demon- strate that the central axiom of Homo economicus is refuted. Her basic premise involves the use of multiple caretakers in human groups to ameliorate the costs of large. extremely diverse geographic distribution and ecological patterns (e. 2005) effectively argues for envisioning humans. Fuentes. and human ancestors. posits at a system where more individuals than the mother are integral to the successful raising of young. (2004) suggest that cooperation is the normative mode of interaction and exchange between human groups. what are the characteristics that define this uniquely successful system in Homo today ? In a nutshell they are: hyper-cooperation at multiple levels. Barton & Hurtado. In fact. For example. patterns of cooperation and social reciprocity were dominant.. 2009). Hrdy (1999. Hrdy. kin selection). heavily dependent offspring with very slow devel- opmental trajectories. Here we see that intergenerational cooperation and cooperation between nonkin.. relatively frequent coor- dinated intergroup violence. Gettler (2010) even pro- vides a strong argument. This does not mean that humans are all egalitarian or selfless. Over the last decade or so a number of researchers have reintroduced a focus on coop- erative interactions as the focal points for evolutionary questions about human behavior. and Indonesia). Dunbar et al. demographic. 2009. as cooperative breeders. however the recent versions of this scenario credit multiple age/sex classes and even non-maternal kin in the group with contributing to the successful rearing. selfishness as a pri- mary pattern was not found in any of the societies studied. Africa. Henrich et al. Hill. Often these individuals are assumed to be related females (see Hawkes et al. So. They review a vari- ety of economic experiments conducted across decades and note that the results show consistent deviations from predictions of the Homo economicus model (that humans will exhibit “narrowly economically self-interested behavior”). such that a consistent selfish behavioral strategy will not be . drawing on ethnographic and evolutionary examples. and defending of young. Rather. 2011).. These researchers undertook a cross-cultural experimental study in 15 small-scale societies scattered across the world (in Latin America. Nowak & Highfield. 2010. Fuentes. apparent in human behavior.g. they found more variation in behavior across societies than had previously been reported. Fehr and Gintis (2007) and Heinrich et al. These foci give us insight into the constituent factors involved in this system. extensive niche construction. and other social variables. thus suggesting that what we are looking for may be a conglomerate of behavioral patterns rather than one specific main behavior (Ehrlich. if there is a broader pattern. symbol use and hyper-complex information transfer.e. 2009. or non-close kin. that we should broaden this specifically to include male carrying of young as an important pattern in early members of the genus Homo. and evidence. 1999.. with much variation in details across society based on integration into world markets.

what might the process by which such a system evolved look like across the last two .. C o n fli c t. explanations to account for the observed patterns of cooperation across human societies (see http://www. increasingly complex sociality and patterns of intergroup cooperation created a particular niche construction pattern that became a central component of human behavior. Specifically. nor evolutionary models based on kinship or reciprocal altruism are sufficient.hss.g. see also Fuentes. Conflict. In such systems the groups that effectively engage in intra. edu/~jensming/roots-of-sociality/). or effective.caltech. (2010. Bulte. This notion of cooperation as a central systemic aspect in human evolution is summa- rized and supported via mathematical modeling by Nowak in his “supercooperators” the- ory (see Nowak & Highfield. arguing that it was the use of long-distance trade routes that enabled humans to out-compete Neanderthals. 2004) and Hart and Sussman (2008) also make the case that cooperative systems were central to the success of earlier hominins and especially the first members of our own genus (e. Fuentes et al. Looking at the fossil record.and intergroup cooperation have higher potential success than those that do not emphasize cooperation or than those that fail to achieve sufficiently effective group-wide or population-wide coopera- tive patterns. socio-political regulation. From these results and a survey of the broader literature they argue that both experimental and field data from across the social sciences demonstrate that neither assumptions of narrow economic self-interest. Evolution of the Human System: Cooperation. and/or competition). Basically. he illustrates via multiple lines of modeling evidence that it is the ability and practice of intense cooperative behavior and patterns of cooperative relationships amongst group members that creates the context for a long-term selective environment (a social and ecological niche) in which the cooperative patterns are a central driver of evolutionary success. these authors explore the success of Late Pleistocene Homo sapiens groups over their contemporaries and the modern day endurance of ethnic traders’ networks in east Africa. and Niche Construction in the Genus Homo If extensive and substantial cooperation is indeed a core part of the process in human evo- lution. and Shogren (2005) extend this con- cept closer in evolutionary time. 2011). Homo erectus/ ergaster). Comparing and contrasting the effects of feedback from cooperation and its concomitant niche construction on the evolution of two different socio-economic processes. These researchers suggest that the selective pressures placed by high predation on early humans had significant effects on their behavior. a n d Ni c h e C o n st ru c t i o n 87 sustainable in human groups. C o o p er at i o n . Oka and Fuentes (2011) and Horan. The main conclusion is that the feedback from intragroup and intergroup cooperation through social networks cre- ates a ratcheting effect on the local ecologies of all groups/organizations responding to similar pressures (predation pressure.

The fossil and material records suggest that the following patterns emerged in the genus Homo between 1.000 years ago) extended period of. and allowed for an expansion of the types and patterns of habitats exploited. In this context Homo experienced increased opportunity for social interactions. combat. range exploration. via feedback systems. including cooperation/reciprocity and conflict – Small-scale trade networks – Geographic expansion • Phase II ~300KYA–40KYA – Neurological reconfiguration – Increased complexity in social identities and social roles – Harsh-ecologies colonization – Expanded Trade networks and material exchanges – Increased variability in intergroup and inter-regional relationships • Phase III ~40KYA–recent – Increased niche variability and ecological patterns – Material inequity and trade expansion/restriction – Symbolic-social niche construction – Increased male aggression and emergence of complex violence– Large-scale regional polities and emergence of complex peace .1 Core Patterns in the Genus Homo • Phase I: ~2–. and effort in.. 2010. 2009.88 Eco l o gi c a l a n d Evo lu t i o na ry Mo d el s million years? What would a pattern of niche construction that gives rise to such “super- cooperators” look like? To assess these questions we need to outline specifically what is proposed to be hap- pening in populations of the genus Homo in the Pleistocene (Box 5. 2011): Increasing brain size (through ~300. or otherwise inhibit the success of pre- dation pressures. reducing the overall selective pressure of predation (Fuentes et al. Hart & Sussman. and testing a variety of novel foraging oppor- tunities. child care.000 years ago (see expanded treatments in Fuentes. increased effectiveness at avoiding predation. increased cooperative interactions between group members across generations.8 million and ~60.1). members of the genus Homo became more costly and predators shifted emphasis to easier prey. an emerging higher cognitive B OX 5. increased communicative complexity.3 MYA – Expanding neocortex – Intragroup Cooperation and social reciprocity central (Inter. Due to increasing abilities to avoid. 2008).and intrasexual and generational) – Increase in material exploitation/manipulation (some specialization/division of labor) – Variable intergroup relationships. These patterns facilitated.

especially the ubiquity of cooperation and com- petition within the social group (Herrmann. In the hominins. patterns of habitat exploitation (e. Silk. In this scenario pri- mates are taking the basal complex sociality of mammals and enhancing it by using social networks/contexts as a tool to meet and modify the demands of the environment (the selective landscape).. becomes more refined in the course of hominin evolution. which is increased in the Miocene hominoids and exponentially enhanced in hominins (see Malone. 2007). 2010). which facilitate diversified. C o n fli c t. all of which creates the ongoing feedback in multiple somatic and extrasomatic systems that facilitate multifaceted engagement and success in a diverse array of environments for the genus Homo. thus changing the selective landscapes for the primate popula- tions. 2012. social niches are established in which selection favors gene complexes/physiologies that predispose or facilitate increased cooperation and com- munication abilities. As the local social and biotic environments are being modified selec- tion pressures are altered. Gowlett. This pattern creates a constant feedback between the social and biotic ecologies resulting in niche construction and the continuous modification of selective landscapes. 2007). This series of interconnected processes is tied to an evolving hominin cognition. Many central components of primate cognition emerged in response to the chal- lenging demands of a complex social life. There is a ratcheting up of this social complexity in anthropoid primates. Hernandez-Lloreda. Call. 2010.g. and trade are inherited and shared among members of groups and local populations. multi-party social negotiations. Fuentes & White. It is at this point in our evolutionary . This extends also to the knowledge and use of local areas. 2011). Increasing cognitive complexity in the later Miocene hominoids begins to facilitate more intensive use of the social relationships as tools to meet ecological challenges. and reciprocity are the primary avenues for social and reproductive suc- cess (Dunbar et al. and increasingly plastic. 2007. Nowak & Highfield. C o o p er at i o n . MacKinnon & Fuentes. and emerges in its modern form via the more recent evolution of the genus Homo (MacKinnon & Fuentes. and thus increased cooperation and reciprocity become central components of behavioral rep- ertoires. Given these patterns and potential processes we can envision a scenario of integrated cooperation as a component of the human niche with central roles for ecological and social inheritance. As environmental challenges also are increasingly negotiated with information transfer via complex communication patterns. In this con- text specific heritable components of human niche construction play core roles. & Tomasello. physiological and behavioral adaptations emerge to effectively negotiate the increasingly complex and information-rich social networks where coalitions.. 2011). cross-generational cooperation. and a diverse pattern of intergroup cooperation and conflict across the human species. Hare. Social tra- ditions such as tool use. Primates are characterized by a specific type of “social intelligence” (Dunbar & Shultz. a n d Ni c h e C o n st ru c t i o n 89 functioning via niche construction and its interplay with selection pressures. The use of fire and increased infant survivorship add to the niche constructing capacity via increased capabilities of extra-somatic modification and new demographic structures. manufacture. 2011). the emergence of symbolic identities. This pat- tern begins within the comparative context of primate evolution.

technological. whereas previously most interfaces were at small-scale intergroup and interindividual levels. While there are countless definitions for “war. As noted above.000 years ago or so. and pat- terns of broad-scale political peace and organized warfare/large-scale lethal aggression in humans. It is in our recent evolutionary history that the current versions of human relatively frequent large-scale coordinated lethal aggression (i. In the earlier phases of human evolution the types and patterns of relationships between groups were different than those in humans of the last 20. and increased scale of conflict and cooperation. emerging in this most recent phase of our evolution. The expansion of human interaction patterns to include larger groups and multiple populations with com- plex social and economic hierarchies can lead to different patterns and structures of niche construction. complex inter. and diverse hierarchical social structures change the ways in which basal patterns of cooperation and competition occur and are perceived by human populations. This is followed by (and facilitates) a geographic range expansion and increases in tool use/creation and possibly much broader incorporation of extra-somatic materials such as fire. There is significant contention about the appearance. It is important to recognize that these patterns. this is the first time in human evolutionary history that we see frequent (relative to the previous two million years of human evolution) coordinated intergroup violence. longevity. are not rooted deep in our evolutionary past but rather are capacities facilitated by the changing demographic. aside . 2004). This period could be seen as the point of emergence of true language. However. Finally.and intragroup relationships. as a part of the niche we continue to inhabit and alter. social. war) and peace emerge. extended allocare and a shift to more complex manipulation and use of the environmental characterizes the genus Homo. and selective processes catalyze the move toward modern human intense reliance on trade.. it is nearly impossible to test such an assertion using material remains in the fossil record. starting at some point in the early Pleistocene. This in turn facilitates increased possibilities for differential roles and pat- terns of exchange within and between social groups impacting the tenor and structure of cooperation and competition. and structural realities of human popula- tions.e. context. In this more recent period of human existence.90 Eco l o gi c a l a n d Evo lu t i o na ry Mo d el s histories that we can begin to see the emergence of unique human characteristics related to conflict and cooperation that eventually involve the use of symbols and language. By this definition.” Ferguson (2008) provides a basic and useful one: organized lethal violence by members of one group against members of another group. coordination at larger societal levels begins to be a dominant mode of interaction. These social and ecological niches facilitate the emergence of increased sym- bolic and informationally dense communication within and between groups and a signifi- cant role for intergroup trade and cooperation (see also Potts. War and Peace The integration of symbolic identities. the impacts of the previous patterns on ecological. multipopulation polities.

Haas & Piscitelli. 2003. Also. but cooperative interactions play a central role. most archeological records show some examples of individuals having died at the hands of others (Walker. engaging in war cannot be viewed as sparking the origin and evolution of cooperation. organized. and lethal with regularity. In addition to iden- tifying specific adaptation of traits or individual behaviors. and many of these can be attributed to cannibalism (either within group or between group) or generalized injuries from interindividual aggression (fights) (Anton. drivers in the evolution of behavioral systems. our attempts to understand the evolutionary histories of humans must include a systems approach wherein liner models of optimality and traditional presentations of natural selection are not the only processes of relevance. ant colonies fight over space. Tattersal. 2011. While warfare is relatively common in recent human history there is very little evidence that wide-scale lethal violence by one group against another characterizes any of the earlier phases of human evolutionary history (Ferguson. warfare is a relatively modern human behavior (10–12. War makes use of the human capacity to cooperate but as complex cooperation predates war by many.000 years ago one finds only a few examples of possible death due to the hand of a conspecific. C o n fli c t. Including the concepts of niche construction and multiple modes of inheri- tance and the realization that plasticity can be seen as an adaptive pattern in and of itself can enable us to make greater headway in assessing and describing the systems involved in human evolution. 2008). Kimble & Delezene. Thus to seriously investigate war as a major aspect of modern humans we need to consider factors such as ecological and social contexts. 2011.1—phase 3) populations of the genus Homo. and many other types of animal groups engage in conflicts. Ferguson. 2001).. 2001). group size and population densities. Clear evidence of actual deaths from weaponry (i. it is interesting to note that large-scale coordinated conflicts such as war require that intensive patterns of cooperation are already in place in the society (Ferguson.000 years old). not specific adaptations. spears and bows and arrows) is only found in modern humans by 8–12.000 years ago. and the dif- ferences in conflict versus homicide versus war. or only. Fry. This defi- nition also distinguishes war from homicide (single events where an individual is killed by another) (Ferguson. From this point on. but are not neces- sarily the main. but none of them are planned.e. many millennia. The human fossil record supports the hypothesis that while some violence between individuals obviously happened in the past. Groups of monkeys may fight over food sources. Conclusion The evolution of human behavioral patterns is complex. Walker. 2009. 2006. Chapter 10. this specific behavior (war) is unique to recent (see Box 5. In fact. 2011). we also need to think in terms of human systems and niches. a n d Ni c h e C o n st ru c t i o n 91 from some populations of chimpanzees. C o o p er at i o n . 1997). . reviewing the available data from the fossil record of humans and other hominins from ~6 million years ago through about 12. Conflict and competition are ubiquitous in living things.

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