Welcome to Territoriality Week! Every day this week, I'll have a post about some aspect of animal or human territoriality. How do animals mark and control their territories? What determines the size or shape of an animal's territory? What can an animal's territory tell us about neuroanatomy? Today, I begin by asking two questions: first, what is the functional purpose of establishing territories? Second, to what extent can we apply findings from research on animal territorial behavior to understanding human territorial behavior?
It seems that everyone becomes an amateur animal behaviorist while walking their dogs. They notice that their dogs tend to pee on - well - just about everything, and infer that Fido is marking his territory. That most people are familiar with at least the basic principles of animal territoriality would suggest that the study of animal territoriality is fairly well established. Indeed, behavioral biologists and ethologists have been interested in animal territoriality since at least the 1920s. The main purpose of animal territoriality, it would appear, is excluding others from certain geographical areas through the use of auditory, visual or olfactory signals or by the threat of aggression. While there are certainly variations, territoriality seems to exist throughout the vertebrate phylum. While many of the early studies of territoriality focused on birds, later researchers investigated territorial behaviors in fish, rodents, reptiles, ungulates (hoofed animals, like cows), and primates. Territories may be held by individuals, by pairs, or by groups. They may be defended against anyone, against only members of the same species, or against only members of the same sex.
Why would territoriality be so widespread in the animal kingdom (at least among vertebrates)?
Dozens of reasons have been offered, including increasing security and defense, reducing the spread of disease, reinforcing dominance structures, and even localizing waste disposal. But an English zoologist named Vero Copner Wynne-Edwards suggested that territoriality operates in order to control population size. Julian Edney, a psychologist from Arizona State University, described Wynne-Edwards's hypothesis in this way:
[T]erritoriality is a link between social behavior (competition and dominance) and population control in many animals. Communities regulate their own numbers by the use of "conventionalized" competition, usually among males, for territory and the accompanying rights to food and (sometimes) mates. The winners are dominant animals and acquire social status, but since they are a fraction of the population, only a few community members get access to space, scarce resources, and females, thus limiting the size of the next generation. The next generation is also guaranteed food, because winners of territory spread themselves thinly over the terrain. Thus the habitat's food sources are not exploited beyond regenerative capacity, and a reasonable supply is ensured for the future.
In other words, the size of the population, and therefore the availability of resources for individuals within the population, is controlled by virtue of the fact that territory winners are generally the only lucky individuals who get to breed and pass their genes on to subsequent generations. Edney notes that Wynne-Edwards's theory is particularly attractive because, at least on some level, it applies to humans as well. It isn't much of a stretch to note that there is an observable relationship between territory ownership and social status, or between territory size and social status, in humans. For example, for much of modern history, one had to be a land-owner (not to mention white and male) in order to participate in government or even to vote. The corner office is so highly prized in business buildings partly because it is bigger than other offices.
It should come as no surprise to the frequent reader of this blog that I would argue that since humans are just another species from among many, a theory regarding animal behavior more likely than not ought to apply to human behavior as well.
I find Edney's description of the origins of human territorial behavior quite interesting, especially considering the historical context in which he was writing. While he seems content to use animal behavior as an analogy for human behavior, Edney is quick to note that territorial behavior in humans, while similar in appearance to animal territoriality, may have different origins. He objects to the practice by which some other researchers would "beastopomorphize" humans. He writes,
...it does not follow from the gross similarities between the territorial behavior of some animals and man that the underlying mechanisms are the same in both, nor that they are genetic. To assume so, incidentally, has an interesting political consequence: It relieves man of the moral responsibility for his territorially aggressive acts and invites the rationalization of human territorial warfare as simple fulfillment of man's genetic predispositions.
He offers the following as evidence that human territoriality is different from animal territoriality, and in particular, is not derived from biology:
- (a) Human use of space is very variable and not like the stereotypic spatial expressions of animals. This suggests a learned, rather than a genetic, basis.
- (b) The association between territory and aggression, treated as fundamental by [some researchers], is not clear-cut in humans.
- (c) Territories serve primarily "biological" needs for animals (shelter, food sources), whereas humans use them also for secondary purposes (e.g., recreation).
- (d) Animals usually use only one territory and for continuous periods of time. Humans may maintain several territories (home, office, mountain cabin) in different locations.
- (e) Humans also "time share" temporary territories (e.g., tables at a restaurant), whereas this is rare among animals.
- (f) Total invasion of one group's territory by another is rare among nonhuman animals but occurs in human warfare.
- (g) By virtue of their weapons, humans are the only organisms that can engage in territorial warfare without trespassing.
- (h) Humans are also the only territorial organisms that routinely entertain conspecifics on home ground without antagonism (as in visiting).
Have the distinctions between humans and animals that Edney laid out (above) in 1974 held up in the face of empirical research? Do you think that human territoriality is qualitatively distinct from animal territoriality, or only quantitatively so? Do you think that human territoriality is purely the result of learning, experience, and/or culture? Or is human territorial behavior built upon evolutionarily ancient mechanisms, subsequently modified or shaped by culture?
Please jump in with your thoughts in the comments! Subsequent posts this week will address some of these, and other, questions about territorial behavior in humans and non-human animals.
Edney, J. (1974). Human territoriality. Psychological Bulletin, 81 (12), 959-975 DOI: 10.1037/h0037444