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
Dog statue image via Flickr/THEfunkyman. Whitehall Estate image via Flickr/Steven_M61.
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So, what does it mean that the men working on our office building have taken to using the ladies room and leaving urine on the toilet with the seat up, despite the presences of perfectly adequate mens rooms at no further distance. Are they marking our territory as theirs or trying send a sexually aggressive message?
Here's one from the rural outreaches of N. Carolina. The coyotes have started leaving scat in my dog's territory and visa versa.
I want to keep my dog. I don't want to kill coyotes. What to do???
Tracy, I think they just like the smell of your poo.
I think a phenomenon that is so universal, as is human territorial behavior, must have some biological basis. Probably the tendency to group identification, which seems to be natural in humans, accounts for this universal occurrence of territoriality. It can be noted that one example of human territory sharing is in the case of the Australian Aboriginal people. Even though they had quite well defined lands, they could share their territory with other tribes, who's lands had been affected by drought. Human territoriality may be natural to some degree but it is also clearly culturally quite variable.
Territory marking also serves to identify members of a pack, and there-by identify intruders as well. Dogs in a given area form distributed packs based upon barking behaviors, as well as familiar scat.
Years ago one of my dogs led me to understand the barking chaos that accompanied every intruder (mailmen, UPS trucks, etc)...and why dogs continued to bark when the intruder was clearly no longer an immediate threat.
The code turned out to be beautifully simple: barking intensity/frequency defines both the location of the intruder and a safety zone for those unable to defend themselves...with continuous updates till the intruder has left pack territory. The value of pack behavior that warns intruders of a defending army, and provides safety information is immeasurable.
For cats, the chin-rubbing scent marking is a way of knowing where they've been. Each time I've moved with cats, I've scratched under their chins then methodically marked tracks for them - out the window, down the banister, down the stairs, and out along the edges of the yard in sheltered paths -- before letting them out the first time.
It's quite clear they're following those tracks, sniffing and renewing them. Once they venture away from the lines I marked I can see a much more tentative progress, more looking around and up -- and when they get startled they run until they cross one of those lines then turn and follow it back to safety.
The same scent marking works for new scratching posts
I can also see the neighbor cats that come to the same space from elsewhere notice the scent marks too.
Result, on a sunny day, a 'clowder of cats' -- sometimes half a dozen or more scattered around the sunny warm yard, each carefully far enough from the others that they can all relax for a while.
It sounds pretty far fetched to me that human territoriality would have absolutely no biological basis. Since humans are socials creatures, wouldn't it be best to compare territoriality on the group level rather than the individual level? Or perhaps group territoriality has a larger biological basis than individual territoriality does. Also, some non-human animal territories temporarily dissolve under high resource availability, so maybe the issue would be best investigated related to some limited resource.
e) Timeshare. The various pods of Orca will time share a single unique pebble scratching beach on the East side of Vancouver Island. They are polite to awaiting pods, and appear to share gratefully what looks like a pleasurable activity.
@Tracey: Not sure what to tell you, though I'm not sure science can help you in this case :-)
@Richard: Fantastic. I was trying to think of an example of this sort of "timeshare" behavior in the animal kingdom, but couldn't come up with anything. Do you know if this behavior has been described in a journal article?
A discussÃ£o de territÃ³rio e seus correlatos tem evoluido nos Ãºltimos anos nas ciencias humanas, sobretudo na Geografia. Acredito que a teoria utilizada estÃ¡ equivocada, porque na atualidade se discute territÃ³rio nas multiplas dimensÃµes, ou seja, polÃtica, economica, cultural e natural.
Cats often have timesharing of territory:
Men smoke cigars to mark territory. That's why it often seems presumptuous or pretentious when a younger man smokes a cigar; it's as if he hasn't earned the status to warrant marking territory.
That's how I think of cigar smokers.
a) Marriage customs and standards of handsomeness vary wildly but that doesn't mean that sex and mating aren't human instincts. Cats' use of territory is variable, too.
b) Visceral reactions, from "get off my lawn" fury and No Hunting to love of the ancestral farm to whatever drives amti-immigrant discrimination, suggest that there are very basic feelings involved.
c) Did someone fail to notice the invasion of North America by Europeans 500 - 100 years ago? Territories (hunting grounds, farm lands, fishing or timber rights, access to drinking water) are extremely important to survival. You just don't notice because you're in a dominant society and a prosperous time. Indian wars were fought over hunting rights. Non-agressive people, like the Inuit, were pushed to the least hospitable land.
d) animals that use different places, like hippos with their daytime river and nighttime pasture, may well defend more than one territory. So will baboons and other monkey with fruiting trees.
e. Cats time-share their territories, especially urban cats with high population densities; Scientific American had an article about it, years ago. As for time-sharing at restaurants--have you ever watched birds at a birdfeeder? They take turns, flight after flight, until an Important Personage like a jay comes along--then they get out of the way.
f. Jane Goodall observed chimps in genocidal territorial warfare when they wiped out a group that had split away from the main troop.
g. Monkeys throw sticks and possibly stones, They also howl or yell to establish territory by show of strength. A lot of animals make territorial displays at their borders and may not trespass but simply "negotiate" moving the border by making their opponent back up.
h. Cats also tolerate others in their territory if food supplies are good. another flexible behavior; just look at the colonies of feral cats that someone feeds. Anecdotally, some cats visit particular other cats and so have friends. I owned a cat that would bring home hungry young cats and would encourage them to eat from its outdoor bowl with little prrpy noises. Wolves have been described visiting relatives in neighboring territories.
Does anybody know of any single cell organisms that exhibit territoriality?
So could this explain at least in part why some woman tend to get involved with aggressive men, time after time. Despite the fact it leads to abusive, controlling and generally unsatisfying relationships. I realize self esteem, self efficacy and family background play a role as well. I have always been a predominately non-aggressive male, especially as former Marines go. I have done a great deal of self study in psychology, philosophy and the sciences in general. I often find myself in the role of sympathetic ear to female friends over the years. They go on and on about the mental and or physical abuse and infidelity. I proposed my own theory that possibly when distracted and caught up in the selection process at the local bar they were driven subconsciously by an ancient instinct. Possibly they were feeling drawn to aggressive men because 10,000 years ago that kept you and you children safe. Which came first, the abusive male or the woman that rewarded him for it?
Except for the example of orcas mentioned above, it would seem only land animals can mark territory, since it would be pretty difficult to establish turf in water. Thoughts?
Do you think the level of testosterone is related to one's desire to own a big and spacious house? Since testosterone is related with dominance and power.