Female Territoriality Research Articles

Territoriality in females of the slender opossum (Marmosops paulensis) in the Atlantic forest of Brazil

Spatial organization within animal populations is often thought to reflect the outcome of strategies implemented by each individual to enhance its reproductive success and survival (Clutton-Brock 1989). Thus, while females usually focus on the acquisition of food and breeding sites, male dispersion is more often determined by the distribution and availability of females (Clutton-Brock 1989). Due to these factors, intraspecific competition for space could lead to the adoption of a territorial strategy, whenever the benefits of territorial defence are higher than the costs (Brown & Orians 1970). Among small mammals, two main hypotheses have been proposed to explain the occurrence of territoriality in females. Ostfeld (1990) proposed that females should defend food resources, so the distribution and availability of food items should determine the cost–benefit relationship of adopting a territorial strategy. However, Wolff (1993) developed a hypothesis, based on small rodents, that females should defend nest sites in order to avoid infanticide, the so-called pup-defence hypothesis.

Home range and habitat preference of female lions (Panthera leo persica) in Gir forests, India

The social organization of Asiatic lions (Panthera leo persica) differs from African lions (P. l. leo) in that breeding lionesses defend resource based territories while male coalitions maximize coverage of female groups. Thus, lion density in the Gir forests of India is dictated by female territory size. We studied the home range and habitat preference of lions using radio telemetry on seven lionesses spaced throughout the Gir between 2002 and 2005. Radio locations obtained by homing in were plotted on a classified (LISS III FCC) habitat map of Gir to obtain habitat use and availability. Habitat preference was computed using compositional analysis and Ivlev’s index. Average (±SE) 100% Minimum Convex Polygon (MCP) range of six lionesses was 48.2 ± 10.6 km2, 95% MCP was 34.7 ± 7.8 km2 and 95% fixed kernel range size was 32.5 ± 8.2 km2. Breeding female group density and group size was about 3 per 100 km2 and 1.3 (0.5 SD, n = 45) respectively. Lions were observed to show a habitat preference ( $$ \chi_{(6df)}^{2} = 11. 4 $$ , P = 0.08), the order of preference was Moist Mixed forests > Mixed forests > Savanna habitats > Teak-Acacia-Zizyphus-Anogeissus forests > Acacia-Lannea-Boswellia forests > Thorn and Scrub forests > Agriculture areas. Habitat preference during the day was for dense vegetation ( $$ \chi_{(6df)}^{2} = 35 $$ , P < 0.001). At night lions even ventured into agricultural fields. Our data suggests that dense habitats are preferred by lions in Gir to escape the heat of the day and to be in good cover when human activity was likely to be at its peak within forested areas.

Female territoriality in a paternal mouthbrooding cardinalfish to avoid predation against spawned eggs

Generally, paternal mouthbrooding cardinalfishes are characteristic of sex-role-reversed animals: females have a higher potential reproductive rate and are more active in mating competition than males, and the operational sex ratio (OSR) is female-biased. However, one species of cardinalfish, Apogon notatus (Houttuyn, 1782), shows unusual sex roles: females alone defend their breeding territories to form pairs, even though the OSR is male-biased. This is inconsistent with the general rule that breeding territoriality is shown by the more abundant sex. We examined the function of female breeding territory in this fish using field observations. Prior to the breeding season, large females established their territories earlier than small females. Earlier settlers occupied deeper areas with larger boulders where conspecifics were less likely to aggregate. As the level of conspecific aggregation increased, spawning females suffered from frequent intraspecific interference and subsequent egg predation, leading to increased time or energy spent on territorial defense. For the females, territories that have more boulders and fewer conspecifics might be of higher quality because such places are safe from egg predation and less costly to defend. We conclude that females defend their breeding territories to avoid predation of spawned eggs rather than to guard high-quality mates or to increase mating opportunities.

Carotenoid-based status signaling by females in the tropical streak-backed oriole

In many tropical bird species, both males and females maintain elaborate plumage traits. Although there is considerable evidence that many male plumage traits function as status signals that convey information about fighting ability, less is known about status signaling in females. We tested whether the carotenoid-based orange breast coloration of the female streak-backed oriole (Icterus pustulatus pustulatus) signals status during territorial interactions. To do this, we simulated territorial intrusions using taxidermic models and compared the roles of the sexes within pairs during territorial defense directed toward different types of simulated intruders. Females were more territorial than their mates during the breeding season, whereas males were more territorial than their mates in the nonbreeding season, contrary to patterns seen in studies of temperate zone birds. The coloration of simulated female intruders also influenced territorial responses: When presented with color-augmented female models, females responded with greater intensity than their mates, whereas the intensity of defense was similar for both sexes when presented with average-colored female models. The greater female response to more colorful intruders suggests that females perceive more ornamented females as greater threats to their territorial tenure or to their pair bond. These results are consistent with the hypothesis that female carotenoid-based coloration signals status in this species. Key words: elaborate monomorphism, female ornamentation, female territoriality, sexual selection, social selection, status signal. [Behav Ecol]

Female defense polygyny in the territorial triggerfish Sufflamen chrysopterum

The occurrence of polygyny requires specific environmental conditions such as female aggregation or patchy resource distribution. However, it is difficult to determine the factors responsible for polygyny in species in which the territories of both sexes overlap. To overcome this, we performed female removal experiments in the polygynous triggerfish Sufflamen chrysopterum (Balistidae) in the Okinawa coral reef. Both sexes defended their territories exclusively against consexuals of the same species, and female aggregation was absent. Each male territory included 1–3 female territories, and nonterritorial males were significantly smaller than territorial males. Further, the body size of territorial males was positively correlated with that of the largest female in their territories, and larger males tended to mate with more females. The results of the female removal experiments (n = 10 females) indicated that females competed for better territories rather than larger mates. In contrast, males abandoned the territories once the females emigrated. These results strongly suggest that males defend females rather than sites and compete for larger and a greater numbers of females. Thus, in S. chrysopterum, female defense polygyny occurs in the absence of female aggregation.

Obituary

Obituary

The function of breeding season chip calls by female hooded warblers (Wilsonia citrina)

We document use of 'chip' vocalizations by breeding female hooded warblers (Wilsonia citrina) and propose explanations for the function of the calls. Fertile females (building or laying stage) chip differently than incubating females, via a lower chip rate, shorter bouts of chipping and more bouts per hour. Females responded to playbacks of fertile chipping and a female model on their territories by approaching and chip calling. Resident males were generally not responsive to their mate's chip calls and did not move closer when their mates chipped; however, neighbouring males often intruded onto territories in response to female chip calls. Neighbouring males did not distinguish between playbacks of fertile and incubation chip calls, but responded to both at equally low frequencies. Radio-tracked neighbouring males intruded onto territories significantly more frequently when females were chipping than when they were silent. Such intrusions likely function as extra-pair mating attempts. Finally, variation in call pattern among fertile females did not correlate with frequency of extra-pair young. Our study provides some support that chip calls of hooded warblers function in female territory defence and attraction of potential extra-pair males but further testing of these hypotheses is required.

Social biology of rodents

Herein, I summarize some basic components of rodent social biology. The material in this paper is summarized and condensed from a recent book "Rodent Societies: An Ecological and Evolutionary Perspective" edited by J. O. Wolff and P. W. Sherman (2007). I describe the four basic spacing patterns and illustrate how female territoriality is a function of offspring defense and male mating tactics are a function of female defensibility. The vulnerability of young to infanticide shapes female spacing and mating behavior. Food does not appear to be a defensible resource for rodents, except for those species that larder hoard nonperishable items such as seeds. Philopatry and the formation of kin groups result in genetic sub-structuring of the population, which in turn affects effective population size and genetic diversity. Dispersal is male biased and typically involves emigration from the maternal site to avoid female relatives and to seek unrelated mates. Scent marking is a major form of communication and is used in reproductive competition and to assess prospective mates, but it is also eavesdropped by predators to locate prey. Females do not appear to alter the sex ratio of litters in response to maternal condition but among arvicoline rodents daughters appear to be favored in spring and sons in autumn. Rodents are relatively monomorphic; however, females tend to be larger than males in the smallest species and smaller in the larger species. Predation risk results from an interaction among foraging time and vulnerability and in turn affects behavioral and life history characteristics.

Behavioral Monogamy and Fruit Aavailability in the Large Treeshrew (Tupaia tana) in Sabah, Malaysia

Behavioral monogamy in mammals varies from male–female pairs that spend most of their time in close spatial contact (associated pair-living) to pairs that occupy exclusive territories but travel, forage, and sleep alone (dispersed pair-living). We present radiotelemetry data on 46 adult large treeshrews (Tupaia tana) from 2 populations in Sabah, Malaysia, that indicate that this species forms dispersed pairs across a range of ecological conditions. Dispersed pair-living was the primary behavioral mating system in primary forest during a major fruit masting event, in nonmasting primary forest, and in selectively logged forest with significantly higher fruitfall than in primary forest. Territories of behavioral partners were spatially concordant, but both male and female territories typically overlapped the boundaries of 1–3 extra-pair territories. Comparison between masting and nonmasting forests indicated that females exhibited better body condition during masting, whereas males exhibited larger home-range areas and longer daily movements. Both males and females exhibited better body condition in selectively logged versus primary forests, but ranging patterns were not significantly different between these habitats. We argue that intraspecific foraging competition is the most likely explanation for the evolution of dispersed pair-living in T. tana.

Linking behavior, life history and food supply with the population dynamics of white‐footed mice (Peromyscus leucopus)

In this paper we review and integrate key aspects of behavioral and life history traits, food supply and population dynamics of the white-footed mouse (Peromyscus leucopus), a species that is abundant and widely distributed across much of eastern North America. Results are based largely on a 33-year mark-and-recapture study in a forest fragment in northwest Ohio, USA. Behavioral plasticity in such reproductive traits as mating system and parental care allows this species to adjust quickly to changing environments. The species has a relatively "fast" life history, with rapid attainment of sexual maturity and high fecundity in the face of high mortality rates. Maximal reproductive effort early in life enables a rapid population response. Food supply, in the form of mast, determines the size of the reproducing population in early spring, which, in turn, influences the size of the late summer peak population. The peak population size is also affected by short-term weather events possibly acting via the food supply. The effects of weather and food on population growth are in part mediated through competition, including defense of space and suppression of reproduction. The inelasticity of female territories appears to set an upper limit to population density.

Seasonal Changes in Home Ranges of the Giant Kangaroo Rat (Dipodomys ingens): a Study of Flexible Social Structure

Knowledge of home-range sizes and the degree of spatial overlap between males and females can help elucidate mammalian mating systems and social organizations. To characterize the social system and mating strategies of an endangered species, the giant kangaroo rat (Dipodomys ingens), we compared home ranges of males and females during the breeding and nonbreeding seasons using radiotelemetry. Boundaries and sizes of home ranges varied seasonally for males, but not for females. During the nonbreeding season, both males and females remained in exclusive territories located in the core of each individual's home range. During the breeding season, home-range size of males increased significantly as mobility of males increased and home ranges expanded to overlap neighboring territories of females. Home ranges also were more uniformly distributed than in the nonbreeding season, and nearest neighbors were significantly more often opposite-sex individuals. Males likely increased home-range sizes to overlap with multiple females and to enhance their opportunities for mating, perhaps by becoming familiar with neighboring females and monitoring those females for signs of receptivity. Although home ranges of females remained similar in size throughout the year, females seemed able to adjust their home ranges in response to neighboring vacancies. We conclude that spacing of D. ingens is flexible in order to meet changing social and environmental conditions.

Song type sharing and territory tenure in eastern song sparrows: implications for the evolution of song repertoires

Song repertoires are thought to have evolved by sexual selection, with larger repertoires being advantageous in both female choice and territory defence. While most hypotheses of repertoire evolution treat different song types as functionally equal, an alternative hypothesis is that song repertoires evolved to allow song sharing with multiple neighbours. In support of this hypothesis, song sparrows, Melospiza melodia, share high proportions of their repertoires with territorial neighbours in at least three west coast populations in North America, and song sharing is correlated with a territorial advantage in at least two of these. We studied song sharing and territory tenure in an east coast population of song sparrows in North America in which song sharing is significantly less common. We found no evidence for a territorial advantage of whole song sharing in our population. We also found no evidence for a territorial advantage for partial song sharing, even though partial song sharing is as common in our population as whole song sharing is in the west coast populations. Population demographics (such as annual survival and territory density) do not seem sufficient to explain different levels of sharing between populations. Thus, we found no evidence in our population to support song sharing as a target of selection in the evolution of song repertoires.

A Feminine "Writing that Conquers": Elizabethan Encounters with the New World

A Feminine "Writing that Conquers": Elizabethan Encounters with the New World Kristen G. Brookes Farmington, Connecticut Introduction A man who represents his nation delights in exploring a virginal body of land, then takes possession of "her" through an act of seizure figured as penetration. This story of colonization is, of course, a familiar one. Yet what happens when, as in Elizabethan England, the invading nation is represented by a woman, and a virgin at that? Clearly, the fact that the prince who might have been urged to seize America's maidenhead was also a queen presents a potentially shocking metaphorical situation, one that most authors and champions of the imperial project would have viewed not only as an embarrassment, but as an abomination.1 Remarkably, however, rather than shunning this sort of metaphor, several Elizabethan writers and artists exploited it, creating grand visions of an empire founded on a relationship between two bodies figured as female virgins: Queen Elizabeth I, identical with her island realm; and the New World, a vast expanse of "virgin" terrain. Two embarkation poems—Stephen Parmenius's "De Navigatione" (1582) and George Chapman's "De Guiana, carmen Epicum" (1596)—along with the "Armada" portrait of Elizabeth, attributed to George Gower (1588), especially capitalized on the possibilities offered by a same-sex imperial encounter, portraying these New World/Old World "twins" as directly engaged with each other, in relationships ranging from "sisterly" to sexual and predatory. In the past two decades, a great deal of revelatory work has been done on gender and sexuality in discourses of discovery and colonization.2 Because of the scholarship of Annette Kolodny, Michel de Certeau, Peter Hulme, Louis Montrose, and others, the colonial encounter has come to be understood as a heterosexual one, emblematized by that between the standing, armored Vespucci and the naked, reclining female America in Theodor Galle's engraving (ca. 1580) as well as by Sir Walter Ralegh's hortatory proclamation that "Guiana is a Countrey [End Page 227] that hath yet her Maydenhead."3 In a highly influential essay that focuses mainly on Ralegh's Discoverie of Guiana, Montrose explores how gendered and sexualized "protocolonialist 'othering' . . . engages, interacts with, and mediates between two distinctive Elizabethan discourses: one, articulating the relationship between Englishmen and Spaniards; the other, articulating the relationship between the woman monarch and her masculine subjects."4 Taking among his points of departure Galle's engraving and Ralegh's proclamation (which serves as his essay's epigraph), Montrose elaborates on Certeau's notion of "writing that conquers," noting that Certeau takes the engraving of Vespucci's discovery of America as "'an inaugural scene'" of "'writing that conquers,'"—that is, a "'colonization of the body by the discourse of power[,] . . . [which] will use the New World as if it were a blank, "savage" page on which Western desire will be written.'"5 Montrose further defines this term as "the writing subject's textualization of the body of the Other, neither as mere description nor as genuine encounter but rather as an act of symbolic violence, mastery, and self-empowerment . . . in particular, as a mode of symbolic action whose agent is gendered masculine and whose object is gendered feminine."6 Montrose's nuanced essay has made invaluable contributions to English Renaissance studies on gender and colonization. Most influential has been his bringing together of the sexualized relationship between male colonizer and female territory with the sexually charged relationship between this same male figure and Queen Elizabeth I, along with his highlighting of the "land-as-body" discourse that identified the body of Elizabeth with her "island" realm, which, as he notes, made possible analogies between Elizabeth and the New World terrain some Englishmen wished to explore.7 Yet Montrose's focus on Ralegh's negotiation of English masculine identity in relation to the Spanish and in relation to his female sovereign excludes the possibility of a scenario in which the masculine plays no role or in which Queen Elizabeth I is a feminine agent of...

Harvesting of males delays female breeding in a socially monogamous mammal; the beaver.

Human exploitation may skew adult sex ratios in vertebrate populations to the extent that males become limiting for normal reproduction. In polygynous ungulates, females delay breeding in heavily harvested populations, but effects are often fairly small. We would expect a stronger effect of male harvesting in species with a monogamous mating system, but no such study has been performed. We analysed the effect of harvesting males on the timing of reproduction in the obligate monogamous beaver (Castor fiber). We found a negative impact of harvesting of adult males on the timing of parturition in female beavers. The proportion of normal breeders sank from over 80%, when no males had been shot in the territories of pregnant females, to under 20%, when three males had been shot. Harvesting of males in monogamous mammals can apparently affect their normal reproductive cycle.

Territoriality, Site Fidelity, and Survivorship of Willow Flycatchers Wintering in Costa Rica

AbstractWe studied wintering Willow Flycatchers (Empidonax traillii) in two seasonal freshwater wetland habitats in northwestern Costa Rica during five boreal winters, to determine habitat occupancy, overwinter and between-year site and territory fidelity, and the degree to which the sexes maintain and defend winter territories. Both males and females used agonistic displays, song, and other vocalizations to maintain and defend mutually exclusive winter territories. Males were generally more abundant than females, but this varied by site and year. There was no significant difference in male and female territory size, nor any indication of sexual habitat segregation. Similarity in morphology and aggressiveness between the sexes may account for the lack of habitat segregation and the ability of females to maintain territories at wintering sites. Each year, 80%–92% of banded flycatchers that were present in midwinter remained at the site until late winter; of these, 86%–100% of individuals maintained the same territories throughout the entire period. We also observed nonterritorial floaters that subsequently established and held winter territories. Between-year site fidelity averaged 68%, and almost all returning birds established territories with boundaries similar to the previous year. Between-year apparent survivorship estimates ranged annually from 54%–72%, with no difference between sites but weak support for higher survivorship of males compared to females. Values for winter site and territory fidelity were generally higher than those reported for other species and for Willow Flycatchers on the breeding grounds; between-year survivorship estimates were similar to those reported for breeding flycatchers.

The Deer Hunter : essai de topologie imaginaire américaine

This essay centers on Michael Cimino’s film on the Vietnam War to explore how the mythology of space in the American imagination is revisited at the end of the twentieth century. In this film the dichotomies between civilization/the wilderness, male space / female territory, national vs. ethnic identities present in the American imagination from Cooper to the classic western through Washington Irving or Mark Twain, are thoroughly revisited through the tragic counterpointing of a Russian working-class community in a Pennsylvania small town and an Asian heart of darkness. Cimino’s heroes find out that the otherness of evil may turn out to be very familiar--but in the end the lid is back on Pandora’s box.

Genetic analysis of male reproductive success in relation to density in the zebrafish, Danio rerio

BackgroundWe used behavioural and genetic data to investigate the effects of density on male reproductive success in the zebrafish, Danio rerio. Based on previous measurements of aggression and courtship behaviour by territorial males, we predicted that they would sire more offspring than non-territorial males.ResultsMicrosatellite analysis of paternity showed that at low densities territorial males had higher reproductive success than non-territorial males. However, at high density territorial males were no more successful than non-territorials and the sex difference in the opportunity for sexual selection, based on the parameter Imates, was low.ConclusionMale zebrafish exhibit two distinct mating tactics; territoriality and active pursuit of females. Male reproductive success is density dependent and the opportunity for sexual selection appears to be weak in this species.

Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika

We studied foraging site partitioning between the sexes in Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid with harem-polygyny. Females maintained small territories against heterospecific food competitors within large territories of males, foraging exclusively at the inner side of their own territories (foraging areas). Males fed as frequently as females in their own territories, but mostly outside female foraging areas, although they frequently entered female territories and repelled food competitors from the territories. Soon after removal of the resident females, however, harem males, as well as many food competitors, invaded the vacant territories and intensively devoured prey of female foraging areas. This indicates that although female foraging areas appear to contain more food than outside the areas, harem males refrained from foraging there when the resident females were present. We suggest that harem males will attempt to keep female foraging areas in good condition, whereby they may get females to reside in male territories and/or promote female gonadal maturation.

TERRITORIALITY, SITE FIDELITY, AND SURVIVORSHIP OF WILLOW FLYCATCHERS WINTERING IN COSTA RICA

Abstract We studied wintering Willow Flycatchers (Empidonax traillii) in two seasonal freshwater wetland habitats in northwestern Costa Rica during five boreal winters, to determine habitat occupancy, overwinter and between-year site and territory fidelity, and the degree to which the sexes maintain and defend winter territories. Both males and females used agonistic displays, song, and other vocalizations to maintain and defend mutually exclusive winter territories. Males were generally more abundant than females, but this varied by site and year. There was no significant difference in male and female territory size, nor any indication of sexual habitat segregation. Similarity in morphology and aggressiveness between the sexes may account for the lack of habitat segregation and the ability of females to maintain territories at wintering sites. Each year, 80%–92% of banded flycatchers that were present in midwinter remained at the site until late winter; of these, 86%–100% of individuals maintained the sa...

Territory revisits reduce intrusion during spawning trips by female yellowtail damselfish, Microspathodon chrysurus

When territory owners temporarily leave their territories, intrusion often increases. Periodic returns (revisits) may reduce intrusion, but not necessarily, because this tactic might increase total time away from the territory. The effectiveness of revisits also depends on a subsequent delay in the return of intruders. Adult female yellowtail damselfish briefly leave their territories in the early morning every 3 days to spawn. Some females make a single trip; others return to their territory several times before they finish spawning. To determine how much time females spent on revisits and whether it reduced intrusion, we observed revisits by multiple-trip females and compared intrusion on the territories of spawning and experimentally caged single-trip and multiple-trip females. If revisits reduce intrusions, multiple-trip females should have higher intrusions when caged than when spawning. If revisits show adaptive plasticity, caged single-trip females should have lower intrusion rates than caged multiple-trip females. Not counting the first trip to the male's nest and the final return trip, we estimated that spawning multiple-trip females averaged 86% of their time at the male's nest, 11% back in the territory and 3% travelling. When owners were spawning, territories of both single- and multiple-trip fish received about 5–7 min of intrusion. When owners were caged, intrusion increased significantly only on territories of multiple-trip females. We consider this the first direct evidence that revisits during temporary absences reduce intrusion on territories, suggesting that revisiting is a plastic trait that females may vary in response to intruder pressure.