Johnson and Toft (2014) argue that humans possess an evolved psychology of territoriality that generates a range of adaptively conditional behavior. While the conditional logic embedded within this territorial psychology is posited to be a human universal, the resulting motivations and behavior are varied as a consequence of adaptively relevant variation in the socio-ecological environment. Thus, in contrast to Audrey’s “territorial imperative” (1966) humans seem imbued with a “territorial conditionality.”

But just what do we mean by “territoriality”? Two questions are often conflated when considering the existence and design of putative psychological adaptations: What is the adaptive problem or set of problems that “territoriality” is designed to solve? What is the design of adaptations that were sculpted to solve them? There are a number of ways to proceed from these questions in the case of territoriality.

The simplest way is to assume that no special adaptationist argument is necessary; in this case, territory is like any other asset and it poses no adaptive problem. As with any other asset, humans may fight to defend their territory. In this scenario, we need only interrogate the operation of known adaptations for aggression (not territoriality, per se) and ask how this particular asset is likely to alter (if at all) the expression of our evolved calculus for violence. This seems to be Szulga’s preferred approach. Another way to proceed is to assume that territory is not like any other resource, and that it may have posed a unique adaptive problem. Thus, Johnson and Toft argue that territory has a unique cost-benefit ecological profile about which humans are designed to reason adaptively. Yet another possibility is that territory may become special when (and only when?) adaptations for imputing sacred value assign special value to territory (Atran, Axelrod, and Davis 2007; Atran and Ginges 2012). Absent the imputation of sacred value, territory may operate in the mind’s eye just like any other asset. Explanations in this domain require us to understand psychological design for the generation of sacred value, not territoriality, per se.

I don’t believe that the evidence as of yet allows us to differentiate successfully among these alternative explanations for the observed phenomenon of “territoriality.” At least – and I may be mistaken – I have not yet seen hypotheses derived and tested that can adjudicate directly among the competing models of territoriality-as-asset, territoriality-as-adaptation, and territoriality-by-product (i.e. by-product of sacred value systems). The first suggests little more than the straightforward operation of adaptations for aggression; the second, the operation of a unique psychology of territoriality; the third, the operation of sacred value mechanisms that generate the appearance of “territoriality” as a by-product.

Because I believe the weight of evidence supports Johnson and Toft, let me proceed here for the sake of argument by agreeing with Johnson and Toft that there are indeed unique adaptations for territoriality. I will refer to “territoriality” as the collection of psychological adaptations designed to respond to adaptive problems relating to the acquisition, maintenance and defense of territory. This would include everything from psychological design for finding certain landscapes more “instinctively” appealing (S. Kaplan 1987; R. Kaplan, Kaplan, and Brown 1989), to psychological decision rules that regulate the use of violence to acquire or defend territory. In consideration of the latter, the asymmetric war of attrition (Maynard Smith and Price 1973; Maynard Smith and Parker 1976) is often modeled to represent agonistic contests between “residents” and “intruders,” in which the former prevails, on average, in contests against the latter. In these situations, the convention “if resident attack; if intruder, retreat” can arise and become stable. In the language of international relations theory, we would say that this environment is “defense dominant” because it is easier to defend than to attack (Jervis 1978).

As Johnson and Toft point out, the possibility of territorial conflict is intensified when both actors view themselves as “resident” and thus are both motivated to attack and expel the other (as may be the case in western Ukraine). Assuming that no two actors can correctly and simultaneously both claim to be residents (an assumption that can be violated), one simple explanation for this dynamic is that for some individuals in some circumstances, it may have been adaptive to “bluff” one’s way into residency – i.e. to behave as “resident” despite being “intruder.” The operation of this “illusion of residency” (viewing yourself as resident when you are intruder) would likely be facilitated by self-deception (Trivers 2011), in much the same way as overconfidence, or positive illusions (Johnson and Fowler 2011).

What conditions might facilitate such an illusion? As Johnson and Toft rightly point out, our evolved psychology is designed to be adaptive in ancestral environments, not modern ones. One very dramatic difference between ancestral and modern environments is nomadism. Notwithstanding the epic territorial shifts and grabs that characterized the Age of Empire (e.g. the “Scramble for Africa”), the modern international system is characterized by a relatively stable and fixed distribution of territorial nation states. This simple fact has many implications that can deeply effect the costs and benefits of territory and its defense: “running” or “hiding” from attack is increasingly implausible; your neighbors today are your neighbors indefinitely, etc. Alen Grafen (1987) has argued that a very different situation arises when “the winner of a fight is likely to retain the territory for a long time.” Where ownership or residency is relatively stable and new patches of land are few and far between, the Bourgeois strategy may in fact break down (i.e. fail to be an ESS) and it may pay both actors to play Hawk. It is precisely in this environment that positive illusions would be particularly beneficial. This qualification to the asymmetric war of attrition seems to fit the modern international system remarkably well. Although space limits further development here, future research should consider ways to test hypotheses that can adjudicate among explanations for this false residency problem.

Lastly, many have worried that the application of the hawk-dove game in international relations is problematic because it applies a micro-framework to macro-dynamics. This is a valid concern; but again, the need is for clearer questions and testable hypotheses: When do groups behave as individuals (and when don’t they)? How does our unique coalitional psychology interact with the incentive structures provided by modern institutions? This is an exciting new frontier of research (e.g. see Boyer and Petersen 2011; Alexander and Christia 2011), and the sensible caution against deriving macro trends from micro principles should not lead us to falsely conclude that macro dynamics are not built upon micro foundations.


Alexander, Marcus, and Fotini Christia. 2011. “Context Modularity of Human Altruism.” Science 334 (6061): 1392–94.

Ardrey, Robert. 1966. The Territorial Imperative: A Personal Inquiry into the Animal Origins of Property and Nations. BCE edition. New York: Atheneum.

Atran, Scott, Robert Axelrod, and Richard Davis. 2007. “Sacred Barriers to Conflict Resolution.” Science 317 (5841): 1039–40.

Atran, Scott, and Jeremy Ginges. 2012. “Religious and Sacred Imperatives in Human Conflict.” Science 336 (6083): 855–57.

Boyer, Pascal, and Michael Bang Petersen. 2011. “The Naturalness of (many) Social Institutions: Evolved Cognition as Their Foundation.” Journal of Institutional Economics FirstView: 1–25.

Grafen, Alan. 1987. “The Logic of Divisively Asymmetric Contests: Respect for Ownership and the Desperado Effect.” Animal Behaviour 35 (2): 462–67.

Jervis, Robert. 1978. “Cooperation under Security Dilemma.” World Politics 30 (2): 167–214.

Johnson, Dominic D. P., and James H. Fowler. 2011. “The Evolution of Overconfidence.” Nature 477 (7364): 317–20.

Johnson, Dominic D.P., and Monica Duffy Toft. 2014. “Grounds for War: The Evolution of Territorial Conflict.” International Security 38 (3): 7–38.

Kaplan, Rachel, Stephen Kaplan, and Terry Brown. 1989. “Environmental Preference: A Comparison of Four Domains of Predictors.” Environment and Behavior 21 (5): 509–30.

Kaplan, Stephen. 1987. “Aesthetics, Affect, and Cognition Environmental Preference from an Evolutionary Perspective.” Environment and Behavior 19 (1): 3–32.

Smith, J. Maynard, and G. R. Price. 1973. “The Logic of Animal Conflict.” Nature 246 (5427): 15–18.

Smith, John Maynard, and G. A. Parker. 1976. “The Logic of Asymmetric Contests.” Animal Behaviour 24 (1): 159–75.

Trivers, Robert. 2011. The Folly of Fools: The Logic of Deceit and Self-Deception in Human Life. Basic Books.


Published On: May 23, 2014

Anthony Lopez

Anthony Lopez

Anthony C. Lopez received a Ph.D. from Brown University in Political Science and is Assistant Professor of International Relations and Political Psychology at Washington State University. His research investigates war as the product of an evolved coalitional psychology, and examines the relationship between inter-group conflict and intra-group cooperation from an adaptationist perspective. Anthony also received training as a Research Affiliate with the Center for Evolutionary Psychology at the University of California, Santa Barbara.


  • So by “territoriality” they mean that humans have to manage space somehow. Big deal. This could not be more different from actual territoriality in the biological sense–i.e. exclusion and defense. My yard mockingbirds and towhees won’t let conspecifics into the yard at all. In contrast, I can go down to UCR and see hundreds of strangers jammed into big rooms together. In fact, many of them show up in my house (at our annual party). People defend space when they feel they need to, otherwise they love nothing better than to throng the marketplaces.

  • Jean says:

    I appreciate this inquiry Anthony. Might it be appropriate to say that you are advocating a form of delineation for concrete, subtle and causative systems of boundaries? It seems to me that one might benefit from using different signifiers (explicit) to point to phenomena and its particular configuration with depth, width and height. For example, the use of the word ‘depth’ as in ‘depth in a product-line’ is not the same as ‘depth in psychology’. The taxonomy of words is often a source of confusion.

  • tmtyler says:

    Proximity-triggered aggression probably counts as a genetic adaptation promoting territoriality amongst organisms that are not highly-mobile. Additionally, many mammals rely on urination and smell to mark territories. Cats and dogs are familiar examples. Stinky pee (and the ability to detect it) may also have other uses – but I think it would be fair to say that these are genetic adaptations promoting defending territory that go beyond mere aggression.

  • Anthony C. Lopez says:

    @Gene: I don’t think that Johnson and Toft’s argument is that humans “manage space somehow.” The fact that humans don’t behave like mockingbirds is not evidence against territoriality in humans; if there are adaptations for territoriality, they would reflect a strategy that would have been adaptively successful in hominid ancestral environments. Thus, Richerson’s critique is apropos if we are serious about evaluating the existence and design of putative adaptations for territoriality. Not whether humans tolerate crowds or attend your parties.

    @Jean: If you are saying that I am saying that it would be nice to have more clearly stated adaptationist hypotheses that are subject to empirical testing, then yes!

    @tmtyler: Interesting points, thank you for sharing.

  • russell1200 says:

    I recall from a book about cats, comparing their behavior with primates, that primates are territorial because the fruit from a tree that they ate at will eventually grow back, while the mouse will not be reborn to the same location. Thus primates have stronger mapping within their repertoire. Note this has nothing to do with fighting anyone over something, but it explains our interest in geography.

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