Traditionally, the role of exhibiting territorial behavior and defending a territory from the intruder has been ascribed to the males. We will try to test the assumption by reviewing specific territorial patterns of behavior in regard to female dominance. Territorial behavior may manifest itself through scent marking, calls, or displays (Kruuk 1972, 1989; Peters and Mech 1975; Price et al. 1990; Smith 1968) and stem from economical, evolutionarily reasons or the reasons of safety (Lewis and Murray 1993; Maynard Smith 1976). Territorial behavior may be made explicit through direct aggression and the extinction of the trespasser. Females, however, may display specific types of aggression or territorial adaptivity due to gender bias (primary role in bearing and raising offsprings) and social hierarchies where females may occupy dominant positions.
Territory plays an important role in animal societies. As Powell defines it, it is “an area within an animal’s home range over which the animal has exclusive use, or perhaps priority use” (70). As we can see, the key concepts within a definition are the phenomenon of home range (HR) and exclusivity or priority. The home range is usually treated as “the area the animal traverses during a certain time ‘in its normal activities of food gathering, mating and caring for young’” (Burt, 1943: 351, qtd. in Lehmann and Boesch 642). As Moyer proves, the phenomena of territoriality and home range in relating to all the animal species are not identical but inter-corresponding. The scholar terms HR as “the geographical area most frequented by the animal in question” (173). Whereas scholars mostly agree on the HR definitions, there are arguments about territoriality conceptualization.
Both phenomena, however, are empirically manifested through specific animal behavioral patterns. Moyer states that territorial behavior as “territorial defense [which is] central home range fighting” (173). Therefore, one may assume that the key distinction between territorial behaviour and the HR behaviour is the type of stimuli, the degree of aggression or agonism and its direction.
Assuming that HR-contextualized behaviors lie in the core of territorial behaviors, one should categorize the former. As far as home-range models of different specie are concerned, researchers are occupied predominantly with the ecological and social variables with special attention to the HR size. Lehmann and Boesch name five factors to get the better understanding of home-range sizes for definite specie and populations (642). These are: size and distribution of food patches, first; predation pressure, second; competition with neighbors for food, mates, or other important resources, third; group size in the case of social animals, fourth; and the availability and distribution of important resources such as water, nest sites, or sexual partners, finally (Anderson, 1984; Barton et al., 1992; Adams, 2001; McLoughlin and Ferguson, 2000). One may hypothesize consequently that animals exhibit territorial behavior being stimulated by food searching, defense, competition, fitness or mating and multiplying.
All the aforesaid stimuli being derived from HR factors stand either separately or altogether in underlying territorial behaviors within specie. Lehmann and Boesch observe that home range within a species depends on many factors acting altogether as a solid system (Chapman and Chapman, 2000) (642). On the ground of recent studies “on use of space” they predict that simpler and general models of a home range depending on a single factor are likely to be substituted by more complicated systems depending on a system of factors (Gordon, 1997) (ibid.). The argument about multifaceted model of home range in regard to the factors contributing echoes somehow with Moyer’s statement about territoriality being a far less investigated issue than home range of animals.
Whereas in regard to HR researchers are primarily occupied with the conceptualization of factors that make animals establish this or that home range, in regard to territoriality they are interested in chaining specific environmental, social or economic factor and a specific type of behavior. Territorial behavior is “such a diverse phenomenon that a single definition seldom includes all of the behaviors considered relevant by all investigators” (Moyer 170). The wider scientific community agrees on projecting anthropomorphic motivational states (ibid.), or, to put it simply, human-like behavioral patterns, onto the animals.
For the present investigation it is important to summarize the motivational states which contribute to territoriality manifestations among different animal specie. Stevenson-Hamilton observes severe resentment of intruders on the feeding spot by the blue Wildebeest (Gorgon taurinus) (Moyer 171). The researcher draws the causal line between anthropomorphic motivational states (defend, protect, resent, oppose) with the physical consequences of conflict, or fight, and territorial behaviors of the animals. Crook creates a causal model stating that, “In animal societies, individuals do not fight because they have territories, they have territories because, among other things, they fight” (quoted in Moyer 171-2). Thus, one distinctive point about territoriality is the prohibition of trespassing exercised by a territorial individual: “the various animals under study defend their territories and protect them from encroachment by conspecifics, or by male conspecifics, or by any intruder” (Moyer 171).
As it has been stated above, there is a causal relationship between territorial aggression and home range: “the closer [the animals] are to the center of their home range, the more likely they are to react to another animal with threatening gestures and fighting responses” (Moyer 171). By the way, the aforesaid territorial responses are better investigated than territorial “stimulus characteristics” (Moyer 172). Among the latter there are the factors of seasonality and life cycle (Moyer 172) besides the location within the HR. Moyer, therefore, suggests investigating “the physiological state of the animal during that aggressive period” (ibid.), thus, age, maturity and fiteness.
Brown states that animals are territorial only due to a limiting resource which is “a critical resource that is in short supply and limits population growth (Brown 1969)” (Powell 71). The proponents of such an approach are Brown (1969), Fretwell and Lucas (1970), Maynard Smith (1976), Watson and Moss (1970). Ebersole (1980), Hixon (1980), Powers and McKee (1994), Saitoh (1991), and Schoener (1981) target food as the most common limiting resource. However, food territorial stimulus may be supported by territory size or space (Wolff 1989, 1993 suggested looking for some other stimulus besides food even if food limiting resource pops into an eye), though it refers mainly to birds and fish. Yet pine voles (Microtus pinetorum) may behave territorially in regard to their tunnel systems (Powell and Fried 1992); and beavers may fight for dams and lodges (Powell 71).
Carpenter summarizes that territorial behavior may be displayed in various forms and factors; among them there are population pressures, social organization, fluctuations of food supplies, predation, and many other factors. However, the researcher underlines that territorial behavior embraces far more intricate factors of the psychological nature being organized in “complex and dependent behavior systems which are organized upon numerous subsystems and behavioral determinants” (qtd. in Moyer 172). Since mental and neuropsychological organization of animals differs from the ones of humans, certain difficulties may exist with categorization of the aforesaid behavioral systems in regard to animal territory after anthropomorphic models. Nevertheless, researchers prefer to take correlations between habitat factors and territorial aggression after human-like patterns a priori.
Powell is resolute in the assumption that territorial behavior is not restricted to one species and is not universal for all the representatives of this specie (71). Garshelis and Pelton (1980, 1981), Powell et al. (1997), Rogers (1977, 1987) have investigated and proved that within one species different individuals may defend territories in different parts of the home range without any single pattern (Powell 71). That is true in regard to black bears. Brown argues that the underlying of such variability within territorial behavioral models is economical defense (1969). Carpenter and MacMillen (1976) have proposed a model where territorial behavior being exhibited by an animal depends on the productivity of food resources and the resource limits.
When productivity is low, the costs of defending a territory are not returned through exclusive access to the limiting resource. When productivity is high, requirements can be met without exclusive access. The model developed by Carpenter and MacMillen (1976) is broadly applicable because it expresses clearly the limiting conditions required for territoriality to exist and it incorporates limits on territory size from habitat heterogeneity, or patchiness. (Powell 71)
Whilst a species characteristic does not seem to contribute significantly to specific territorial behaviors, various classifications of territories in regard to animals (communal territories, functional territories, etc., Burt) seem to affect territoriality. The latter point makes one think more intensely on the correlation between territoriality and gender. Really, identification of different territorial categories may depend on an animal species in regard to its social hierarchy that includes gender factor. For the present analysis, it is very important to include a gender factor and a concept of female dominance into the model of territorial behavior.
There are male dominance, female dominance, male-female and female-male dominance hierarchies (intrasexual and intersexual) being marked either as strong or loose. The hierarchic pattern seems to be predicated by the productivity, predictability, and patchiness of the limiting resources (Bekoff and Wells 1981; Doncaster and Macdonald 1992; Kruuk and Parish 1982; Macdonald 1981, 1983; Macdonald and Carr 1989; Powell 1989; in Powell 70). For the goals of the present synthetic paper it is important to draw “a behavioral phenomenon called female priority” (Wright 138) into analysis.
Wright reminds that there is a correlation between “strict seasonality of breeding and birthing” (ibid.) and female priority in some animal classes such as the Malagasy or New World primates (Jolly, 1966, 1984; Pollock, 1977; Richard, 1978). He observes that most Malagasy primates live under the laws of strong matriarchate with female individuals having leadership, feeding priority, and dominance. Some specie of New World also demonstrate models where females choose and organize travelling routes and have primary access to feeding spots (Boinski, 1987; Wright, 1984; Milton, 1984) (ibid.). Wright also states that, “female leadership may be related to energetic demands of reproduction (Wright, 1984)” (ibid.) as well as female mate choice and sperm competition. All the behavioural patterns (leadership, choice and competition) imply aggression and priority which is the key characteristic of either territoriality or social hierarchical behaviour. They are caused by anti-predator defense and the breeding social benefits (Janson, 1984) (Wright 138) in case of the larger species of New World primate and in some prosimians (Milton, 1985; Kappeler, 1993).
The vast corpus of research is dedicated to investigating male dominance models in relation to territoriality regardless of the fact that territorial behavior may be exhibited either by an individual, or mated pair, or family. We should like to concentrate, though, on territorial female behaviors which vary from those exhibited by male individuals of the same species. It is due to evolution that in each social group males are assigned to perform territorial and defense roles, while females perform specific roles. Females are responsible for giving birth to offsprings, their lactation (in case of mammals) and education or defense during the certain period (the term of maternal care varies on a species, genus and class). It does not matter that they can not display territorial behavior in that sense that territoriality means aggression to some of the aforesaid factors.
Lehmann and Boesch speak about the socio-ecological model that prescribes the rate of female reproduction being directly correlated to environmental jeopardies and the supply and access of resources (642). Thus, females may share with males such territorial aggression stimuli as food supply and space limiting resources. At the same time, because of high reproductive value female individuals are expected to add to the male-oriented list specific territorial behaviors that are rationalized by gender.
As Engh et al. states, in hierarchically structured animal societies (that is especially true about group-living mammals) “sexual selection has favored males that are larger, stronger, and more pugnacious than females” (Darwin, 1871; in Engh et al. 193). However, in some animal societies sexual roles may be reversed in regard to the functions that are traditionally assigned to male individuals. One of the most evident examples of matriarchic dominant behavior that may be classified as territorial is observed with spotted hyenas (Crocuta crocuta). In social groups of Crocuta crocuta structured in linear dominance hierarchies (Frank, 1986; Holekamp and Smale, 1990, 1993; Kruuk, 1972; Mills, 1990; 193) females dominate both in physical parameters and in sexual roles.
That is, among adult hyenas, females are larger and more aggressive than males, and females dominate males in all dyadic contexts (Frank, 1986; Kruuk, 1972). Furthermore, fighting among adult male hyenas is less frequent and less intense than is that observed among adult females (Frank, 1986; Kruuk, 1972; Mills, 1990), suggesting that male-male contest competition over mates may not function as importantly in sexual selection in hyenas as it does in many other mammals. (Engh et al. 193)
To put it in other words, Spotted hyenas are gregarious clan-organized carnivores. Their adult females grant offspring with the heredity of a social rank. In the clan adult males are immigrants mostly. Territorial female hyenas “always have higher priority of access to food than immigrant males” (Engh et al. 199), first, and are defended against forced copulation by immigrant males due to their original genital morphology (ibid.). It is evident that female Crocuta crocuta would not need to behave territorially in regard to male conspecifics on food supply basis. It is logical to assume that intrasexual aggression is displayed more often than intersexual one. Besides, territorial aggression towards the trespassers from other specie may occur on side of females of the clan.
In regard to the major of species in the Carnivora, researchers (Powell 1979, 1994; Rogers 1977, 1987) observe intrasexual pattern of territoriality. It means that individuals “maintain territories only with regard to members of their own sex” (Powell 73). As it is in case of Crocuta crocuta, these species “exhibit large sexual dimorphism in body size” and their “females undoubtedly selectively polyandrous” (Powell 73). Intrasexual territoriality yields in the severity of manifestation to intersexual territoriality when species gain food supplies through resource depression (i.e., food resources are renewed rapidly, Powell 70).
Intrasexual pattern of territoriality may also be observed in case of many social animals when the hierarchy grounds on the ability to compete for food or mates. Dennehy names several factors contributing to displaying intrasexual aggression among pronghorn females: body mass in reindeer (Hirotani, 1990); age in red deer, Cervus elaphus (Thouless, 1990), bighorn sheep (Festa-Bianchet, 1990), and bison (Rutberg, 1986); and antler size in caribou (Barrette and Vandal, 1986) and mule deer, Odocoileus hemionus (Tomback, et al., 1989) (177). After Byers (1997), as well as Fournier and Festa-Bianchet (1995), Dennehy argues that pronghorn females demonstrate explicitly aggressive behavior in contest for social rank starting from one month old more often than other female ungulates. The researcher argues that such competing pattern is due to loose and non-stable hierarchies which exist among pronghorn groups. By the third month of life female fawns are structured within linear hierarchies and pertain their status (that has been acquired as fawn) life-long (Byers, 1997).
In several ungulate species (African buffalo, Syncerus caffer, Prins, 1989, 1996; bison, Rutberg, 1986; caribou, Barrette and Vandal, 1986; mountain goats, Masteller and Bailey, 1988; and red deer, Appleby, 1980; Thouless, 1990) higher-rank female species receives a privilege in foraging. The researcher has found out that pronghorns, Antilocapra americana, display agonistic behavior between the conspecifics of the same gender while competing for social rank, whilst high-rank and low-rank females stay equal in foraging access.
We may assume that there may be a factor of mating to contribute to female territorial behavior. In species where females are polyandrous (“actually or potentially have multiple mates,” Nunn et al., 646), females are engaged into intrasexual fighting over males. The priority belongs to those of higher quality that is manifested through the so-called exaggerated sexual swellings (Clutton-Brock and Harvey, 1976; Dixson, 1983; Hrdy and Whitten, 1987; Nunn, 1999). The prominent swelling is, by definition, “conspicuous coloration of their perineal region” (Nunn et al., 646) which females of some Old World primates use during the part of their menstrual cycle to demonstrate to the mates when they are most likely to ovulate.
The researchers suggest that female sexual behaviors in mammals may step out as “counterstrategies to male attempts at monopolizing fertile matings and coercing females to mate” (Davies, 1985; Gowaty, 1997; Nunn, 1999; Smuts and Smuts, 1993; van Noordwijk and van Schaik 2000; van Schaik et al., 1999) (ibid.). It is true in case of exaggerated sexual swellings with catarrhine primates: bonobo, Pan paniscus, chimpanzee, P. troglodytes, the gelada, Theropithecus gelada, and the hamadryad baboon (Papio hamadryas), females compete intrasexually for better food supply and better phenotype in reproduction. As Nunn et al. prove that females also face the necessity to make their quality explicit, though some members of academic community think that it is not true. Females really need to fight for males for the goals of reproduction; that is why they need to prove their quality.
One may assume further that besides food and mating limiting resource there is a factor of presence/absence of offsprings that matters for female individuals to display territorial aggression. Powell contradicts Wolff (1993, personal communication) who thought that in regard to the black bears the birthing and breeding assignments play the major role in animal life and, by logic, are the most important stimuli for animals exhibiting territorial behavior (Powell 73). However, Nicholson states that in species with strong female dominance hierarchies (e.g., macaques, baboons), mothers are not in the habit of letting other females or males touch or even approach their infants (34). We may assume that in specie with complex social hierarchies, offspring do play a significant role in shaping female territorial behavior.
Nicholson observes a direct connection between a female’s social rank and maternal behavior as the grounds for territorial aggression. In species with rigid dominance hierarchies there are higher-ranking “laissez faire” mothers (Altmann, 1980) among primates who allow the female conspecifics to touch and care for infants in the course of an adaptive strategy. Then, the independence of infants in their early years may be regarded as positive because females may get pregnant more often and offspring have better chance to survive even without maternal support (Nicolson 34). At the same time, low-ranking mothers in strong female dominance hierarchies may react to aggression on side of other females of the community in forms of grabbing, pulling or approaching by makings infants dependent of themselves on a higher degree. Nicholson observes in species with loose hierarchical relationships (langurs, colobus monkeys [Colobus sp.], howlers) the tendency for mothers letting other females approach infants even in their early days (Hrdy, 1976; McKenna, 1979) (ibid.).
Pride has studied the correlations between optimal group size and seasonal stress within the population of Lemur catta. These diurnal prosimian primates inhabit southern and southwestern Madagascar. As eclectic omnivores (Budnitz and Dainis, 1975; Rasamimanana and Rafidinarivo, 1993; Sauther et al., 1999; Sussman, 1974) are characterized by rigid matriarchate social pattern where females have foraging benefits and are engaged into intrasexual dominance relationships (Jolly, 1966, 1984; Sauther, 1993). They live in matriarchic social groups that guard food resources or territory from trespassers (Sauther et al., 1999)” (550), display targeted aggression depending on season. This species is an excellent sample to measure territorial behavior with regard to female dominance because of its two key features – social female dominance and targeted aggression (Pride 552). Pride has found that cortisol levels corresponded to group composition and size. Females in the group dominate over males and have feeding priority over them. They display aggression only from males, and keep individual space from males. The researcher assumes that “their stresses may be intrasexually induced” (Pride 558).
Blanchard et al. observe the manipulative strategy of female sexually dimorphic polygynous mammals that are reported “to provide a high level of care should bias offspring sex ratio in favor of sons” (The Trivers and Willard model, TWM) ((274). The hint to intrasexual competitive behavior is given in the assumption that “the selective pressure for a female to manipulate offspring sex ratio should depend not only on her own ability to provide maternal care, but also on the quality of other females in the population” (ibid.). In case of marked bighorn sheep, Ovis canadensis, they state that TWM refers exclusively to species where female foraging benefit depends directly on female social position (288). It means that in TWM-affected specie (red deer, reindeer, and horses), dominant females are likely to have forage advantage and produce offsprings with larger mass of the body, whilst in other cases (bighorn and Soay sheep) it may not happen. In the former cases the aggression displayed by females intrasexually is higher than in the latter. But in the former cases females are not able to manipulate sex ratio as freely as in the latter.
To finish, territorial aggression may take different forms depending on social hierarchy. Chapais state that in protective alliances in matrilineal dominance systems dominant females are supported better (Japanese macaques) (201) depending in the body mass of their male and female kin and non-kin allies. Females within such structures may form also conservative alliances where “unrelated high-ranking females were mutually dependent for purposes of maintaining their rank above a subordinate family” (Japanese macaques and vervets) (202). Females may also participate in defensive coalitions, xenophobic and protective alliances at equal pace with males (Cercopithecus aethiops).
To put it into nut-shell, territorial and social behavioral patterns do constitute a very important role in animal societies. Despite common assumption that territoriality, social hierarchy and any forms of aggression are characteristic of males, research has proved that females are also able to display territorial aggression and dominance.
The correlation of territoriality and female dominance is better investigated in social animals. The Malagasy and New World female primates have leadership, feeding priority, and dominance in establishing home ranges and better territories. With the larger species of New World primate and in some prosimians female aggression may be dictated by rank, outer defense (from the predators) and maternal care. In social groups of spotted hyenas, Crocuta crocuta, females dominate males in physical parameters and in sexual roles having better access to resources and spaces. Their territorial behavior is mainly intrasexually oriented. The female ungulates display often intrasexual aggression in ovulation and maternity periods. Some of them (bighorn and Soay sheep) are able to manipulate sex ratio among offsprings regardless of forage access which is an adaptive variation of territoriality.
Researchers take anthropomorphic nature of territorial behavior in animals as a given fact. The territorial patterns exhibited by female primates are often classified in terms being shared with human psychology, sociology and anthropology. Both human and non-human primates are able to demonstrate both intrasexual and intersexual territorial behaviors.
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