Territoriality and Home Range Concepts as Applied to Mammals William Henry Burt Journal of Mammalogy, Vol. 24, No. 3. (Aug., 1943), pp. 346-352. Stable URL: http://links.jstor.org/sici?sici=0022-2372%28194308%2924%3A3%3C346%3ATAHRCA%3E2.0.CO%3B2-O Journal of Mammalogy is currently published by American Society of Mammalogists. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/asm.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Wed Aug 22 16:42:28 2007 346 JOURNAL OF MAMMALOGY TERRITORIALITY AND HOME RASGE COSCEPTS AS APPLIED TO MAhfMALS TERRITORIALITY The behavioristic trait manifested by a display of property ownership-a defense of certain positions or things-reaches its highest development in the human species. Man considers it his inherent right to own property either as an individual or as a member of a society or both. Further, he is ever ready to protect that property against aggressors, even to the extent at times of sacrificing his own life if necessary. That this behavioristic pattern is not peculiar to man, but is a fundamental characteristic of animals in general, has been shown for diverse animal groups. (For an excellent historical account and summary on territoriality, with fairly complete bibliography, the reader is referred to a paper by Mrs. Sice, 1941). It does not necessarily follow that this trait is found in all animals, nor that it is developed to the same degree in those that are known to possess it, but its wide distribution among the verte- brates (see Evans, L. T., 1938, for reptiles), and even in some of the invertebrates, lends support to the theory that it is a basic characteristic of animals and that the potentialities are there whether the particular animal in question displays the characteristic. Heape (1931, p. 74) went so far as to say: "Thus, although the matter is often an intricate one, and the rights of terri- tory somewhat involved, there can, I think, be no question that territorial rights are established rights amongst the majority of species of animals. There can be no doubt that the desire for acquisition of a definite territorial area, the determination to hold it by fighting if necessary, and the recognition of individual as well as tribal territorial rights by others, are dominant characteristics in all animals. In fact, it may be held that the recognition of territorial rights, one of the most significant attributes of civilization, was not evolved by man, but has ever been an inherent factor in the life history of all animals." Undoubtedly significant is the fact that the more we study the detailed be- havior of animals, the larger is the list of kinds known to display some sort of territoriality. There have been many definitions to describe the territory of different animals under varying circumstances. The best and simplest of these, in my mind, is by Xoble (1939); "territory is any defended area." Soble's definition may be modified to fit any special case, yet it is all-inclusive and to the point. Territory should not be confused with "home rangev--an entirely different concept that will be treated more fully later. The territoriality concept is not a new one (see ?;ice, 19-21). It has been only in the last twenty years, however, that it has been developed and brought to the front as an important biological phenomenon in the lower animals. Howard's book "Territory in Bird Life" (1920) stimulated a large group of workers, chiefly in the field of ornithology, and there has hardly been a bird life-history study since that has not touched on this phase of their behavior. BURT--TERRITORIALITY AND HOME RANGE 347 In the field of mammals, much less critical work has been done, but many of the older naturalists certainly were aware of this behavior pattern even though they did not speak of it in modern terms. Hearne (1795) apparently was thinking of property rights (territoriality) when he wrote about the beaver as - HOME RANGE BOUNDARY NEUTRAL AREA ----- TERRITORIAL BOUNDARY NESTING SITE BLANK--UNOCCUPIED SPACE O REFUGE SITE FIG.1. Theoretical quadrat with six occupants of the same species and sex, showing territory and home range concepts as presented in text. follows: "I have seen a large beaver house built in a small island, that had near a dozen houses under one roof; and, two or three of these only excepted, none of them had any communication with each other but by water. As there were beavers enough to inhabit each apartment, it is more than probable that each family knew its own, and always ent'ered at their own door without having any 348 JOURNAL OF MAMMALOGY further connection with their neighbors than a friendly intercourse" (in Morgan, 1868, pp. 308-309). Morgan (op. cit., pp. 134-135), also writing of the beaver, made the following observation; "a beaver family consists of a male and female, and their offspring of the first and second years, or, more properly, under two years old. .When the first litter attains the age of two years, and in the third summer after their birth, they are sent out from the parent lodge." Rlor- gan's observation was later confirmed by Bradt (1938). The works of Seton are replete with instances in the lives of different animals that indicate territorial behavior. In the introduction to his "Lives" Seton (1909) states "In the idea of a home region is the germ of territorial rights." Heape (1931) devotes an entire chapter to "territory." Although he uses the term more loosely than I propose to, (he includes home ranges of individuals and feeding ranges of tribes or colonies of animals), he carries through his work the idea of defense of an area either by an individual or a group of individuals. Sot only this, but he draws heavily on the literature in various fields to support his thesis. Al- though the evidence set forth by Seton, Heape, and other early naturalists is of a general nature, mostly garnered from reports by others, it cannot be brushed aside in a casual manner. The old time naturalists were good observers, and, even though their techniques were not as refined as those of present day biolo- gists, there is much truth in what they wrote. A few fairly recent published observations on specific mammals serve to strengthen many of the general statements made by earlier workers. In speak- ing of the red squirrel (Tamiasciurus), Klugh (1927, p. 28) writes; "The sense of ownership seems to be well developed. Both of the squirrels which have made the maple in my garden their headquarters apparently regarded this tree as their private property, and drove away other squirrels which came into it. It is quite likely that in this case it was not the tree, but the stores that were arranged about it, which they were defending." Clarke (1939) made similar observations on the same species. In raising wild mice of the genus Peromyscus in the laboratory, Dice (1929, p. 124) found that "when mice are placed together for mating or to conserve cage space it sometimes happens that fighting takes place, especially at first, and sometimes a mouse is killed. Xearly always the mouse at home in the cage will attack the presumed intruder." Further on he states, "However, when the young are first born, the male, or any other female in the same cage, is driven out of the nest by the mother, who fiercely protects her young." Similarly, Grange (1932, pp. 4-5) noted that snowshoe hares (Lepus americanus) in captivity '(showed a definite partiality for certain spots and corners to which they became accustomed" and that "the female would not allow the male in her territory (cage) during late pregnancy and the males themselves were quarrelsome during the breeding season." Errington (1939) has found what he terms '(intraspecific strife" in wild musk- rats (Ondatra), RiIuch fighting takes place when marshes become overcrowded, especially in fall and winter during readjustment of populations. '(But when invader meets resident in the tunnel system of one of [the] last lodges to be used in a dry marsh, conflict may be indeed savage." Gordon (1936) observed def- inite territories in the western red squirrels (Tamiasciurus jremonti and T. douglasii) during their food gathering activities. He also performed a neat experiment with marked golden mantled squirrels (Citellus lateralis chysodeirus) by placing an abundance of food at the home of a female. This food supply attracted others of the same species.