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1.
1.  In territorial contests, not only acoustic or other signals, but also the movements of a territorial intruder are likely to influence the response of a resident.
2.  We tested this movement hypothesis by simulating moving vs. stationary intruders into the territories of winter wrens Troglodytes troglodytes , using the same non-interactive song playbacks in both treatments.
3.  Male winter wrens showed a different long-term singing reaction in response to a moving than to a stationary intruder.
4.  One day after experiencing an intruder that was switching between three locations, residents started to sing earlier before sunrise, and they sang more and longer songs at dawn than before the intrusion.
5.  Residents receiving the same playback from one location only reacted by starting to sing later relative to sunrise, and by singing fewer and shorter songs than before the intrusion.
6.  We could not discriminate between the treatments when examining the short-term singing reactions during and immediately after the playbacks. However, our results clearly demonstrate an effect of the spatial behaviour of territorial intruders on the long-term territory defence of residents at dawn, about 24 h after an intrusion.
7.  We argue that spatial behaviour of territorial intruders should be an integral part of the study of animal territory defence behaviour. Investigating long-term changes in territory defence at dawn is a sensitive tool for discriminating between different types of intruders.  相似文献   

2.
Territorial behaviour of a clan of spotted hyaenas, Crocuta crocuta, was investigated in the Kruger National Park over a period of 27 months. These hyaenas were highly territorial, spending ? of their total activities on territory patrol by scent-marking intensively and monitoring 64 marking posts, particularly in border regions. Females, the more philopatric sex, were most active in territory defence. Local intensity of territorial activities by residents within their 130-km2 territory was directly proportional to intrusion pressure by neighbours. When clan size was reduced, the ability to defend disputed land declined and larger neighbouring clans appropriated parts of the territory. We propose a relationship between resource distribution, intrusion pressure and territory defence.  相似文献   

3.
ON THE DEFINITIONS AND FUNCTIONS OF DOMINANCE AND TERRITORIALITY   总被引:4,自引:0,他引:4  
1. Dominance/subordinance is a relationship between two individuals in which one defers to the other in contest situations. Each such relationship represents an adaptive compromise for each individual in which the benefits and costs of giving in or not giving in are compared. Familiar associates in groups or neighbours on nearby territories may develop relatively stable dominant-subordinate relationships based on individual recognition. Although the aggressive aspects of dominance are usually emphasized, the less conspicuous actions of the subordinate individual are actually more important in maintaining a stable relationship. 2. In evolutionary terms, dominance essentially equals priority of access to resources in short supply. Usually the subordinate, who would probably lose in combat anyway, is better off to bide its time until better able to compete at another time or another place. Both individuals save time, energy, and the risk of injury by recognizing and abiding by an established dominant-subordinate relationship. 3. Dominance can be either absolute or predictably reversible in different locations or at different times. Of the various forms of dominance behaviour, rank hierarchies and territoriality represent the two extremes of absolute and relative dominance, respectively. A dominance hierarchy is the sum total of the adaptive compromises made between individuals in an aggregation or organized group. Many animals seem to be capable of both absolute and relative dominance, and within species-specific limits the balance may shift toward one or the other. High density, or a decrease in available resources, favours a shift from relative to absolute dominance. Some species may exhibit both simultaneously. Social mammals may have intra-group hierarchies and reciprocal territoriality between groups, while the males of lek species may exhibit ‘polarized territoriality’ by defending small individual territories, with the most dominant males holding the central territories where most of the mating takes place. 4. Territoriality is a form of space-related dominance. Most biologists agree that its most important function is to provide the territory holder with an assured supply of critical resources. Territoriality is selected for only when the individual's genetic fitness is increased because its increased access to resources outweighs the time, energy, and injury costs of territorial behaviour. 5. Territoriality was first defined narrowly as an area from which conspecifics are excluded by overt defence or advertisement. The definition has been variously expanded to include all more or less exclusive areas without regard to possible defence, and finally to include all areas in which the owner is dominant. I define territory as a fixed portion of an individual's or group's range in which it has priority of access to one or more critical resources over others who have priority elsewhere or at another time. This priority of access must be achieved through social interaction. 6. My definition excludes dominance over individual space and moving resources, and includes areas of exclusive use maintained by mutual avoidance. It differs from most other definitions in its explicit recognition of time as a territorial parameter and its rejection of exclusivity and overt defence as necessary components of territorial behaviour. There is an indivisible continuum of degrees of trespass onto territories, and functionally it is priority of access to resources that is important rather than exclusive occupancy. 7. There is a similarly indivisible continuum in the intensity of behaviour needed to achieve priority of access to resources. Deciding whether or not an exclusive area is defended leads to the pointless exercise of trying to decide which cues indicating the owner's presence are conspicuous enough to merit being called defence. Concentrating on overt defence emphasizes the aggressive aspects of territorial behaviour rather than the equally or more important submissive aspects such as passive avoidance.  相似文献   

4.
In territorial animals that hide to avoid predators, a predatory attack creates a conflict because a hiding animal cannot defend its territory from conspecific intruders. When intruders are persistent, a past conspecific intrusion informs a territorial resident that future intrusions by the same animal are likely. Using a mathematical model, I examine the effects that past territorial intrusions can have on antipredator behaviour. Past territorial intrusions rarely affect a resident animal's time to hide (the optimal behaviour is to hide as soon as the predator initiates its attack). In contrast, past intrusions should shorten the length of time during which territory holders remain in hiding, with the magnitude of this effect depending on the time of the predator's attack, the re-intruder's pattern of return, and the intrusion rates of other conspecifics. The results of the model show that we need more information on patterns of re-intruders' behaviour, and emphasize that a similar functional explanation could underlie other behavioural changes following territorial and/or aggressive encounters (such as winner/loser effects or changes in display frequency and territorial vigilance). Differences between my findings and those from previous studies suggest that the trade-off between antipredator behaviour and territorial defence can involve different costs from the trade-off between antipredator behaviour and foraging.  相似文献   

5.
Some individuals in species with extended periods of territorial occupancy may change territory locations within a single bout of territorial activity. Length of occupancy of mating territories among males in a local population of white-faced dragonflies (Leucorrhinia intacta) varied from more than 6 h to 15 min or less. Males with short tenures often established territories in several locations on the pond during a day. Several hypotheses have been proposed to explain shifting territorial sites rather than remaining in a single site during one bout of territoriality. We attempted to test the hypothesis that males shift to leave low-quality sites. Site quality may be affected by costs of defense in relation to intruder rate and the mating benefits of holding the territory. To test whether variation in these possible effects of benefits and costs of territoriality influenced tenure, we manipulated local quality of oviposition substrate and perch density. The quality of oviposition substrate, but not perch density, influenced both potential benefits and costs of territoriality. Female density was higher in areas with good substrate, but so were rates of males intruding into the territories, rates of chasing by territorial males, and local density of territorial males. More matings occurred in areas with good substrate, but among males with tenures of 15 min or more, mating success per male and tenure lengths did not differ statistically among treatments. Defense costs were low for all treatments and perhaps were not an important influence on tenure duration. Territorial males in this population probably adjusted local density to expected mating success by initial choice of site rather than by varying tenure length. Variation in tenure length at a site resulted, in part, from stochastic external factors, such as predation attempts.  相似文献   

6.
Anthidium manicatum males defend patches of flowering plants as mating territories and copulate with females that enter the territory to obtain nectar or pollen. Territorial males defend their territories not only against conspecific males but also against many other insect species that attempt to utilize the flowers inside the territory. In our study area, a territorial male defended its territory against conspecific males on average only twice per h but it attacked other species of insects on average 70 times per h. Territorial males distinguished between different species of intruders and attacked them unequally. During the hour following experimental removal of the territory holder the number of intruders of some insect species tripled but the number of intruders of those species that were rarely attacked remained approximately the same. A. manicatum males, but not females, bear long spines on the last two segments of the abdomen. Territorial males rammed intruding insects at high speed and shortly before the moment of impact they curved the abdomen forward to hit with the spines. Intruders could be seriously damaged or killed by these attacks. Abdominal spines of A. manicatum males may have evolved as weapons to increase the effectiveness of interspecific territoriality.  相似文献   

7.
Scent marking is ubiquitous among the dwarf antelope and gazelles of Africa, but its function has been the subject of debate. This study examined preorbital gland scent marking in the oribi, Ourebia ourebi, a territorial African antelope. Several hypotheses for the function of scent marking by territorial antelope were tested with observational data. Of these, the hypotheses that scent marking is driven by intrasexual competition between neighbouring males, and that marks serve as an honest advertisement of a male's ability to defend his territory from rivals, were supported best. Thirty-three territorial male oribi on 23 territories marked most at borders shared with other territorial males, and territorial males marked more often at borders shared with multimale groups than at borders shared with a single male. This suggests that males perceived neighbouring male groups as a greater threat to territory ownership than neighbouring males that defended their territories without the aid of adult subordinates. Marking rate was unrelated to territory size or the number of females on adjacent territories, but males with many male neighbours marked at higher rates than those with fewer male neighbours. These results suggest that the presence of male neighbours has a greater effect on the scent marking behaviour of territorial antelope than has been considered previously. Copyright 1999 The Association for the Study of Animal Behaviour.  相似文献   

8.
The availability of food resources can affect the size and shape of territories, as well as the behaviors used to defend territories, in a variety of animal taxa. However, individuals within a population may respond differently to variation in food availability if the benefits of territoriality vary among those individuals. For example, benefits to territoriality may differ for animals of differing sizes, because larger individuals may require greater territory size to acquire required resources, or territorial behavior may differ between the sexes if males and females defend different resources in their territories. In this study, we tested whether arthropod abundance and biomass were associated with natural variation in territory size and defense in insectivorous green anole lizards, Anolis carolinensis. Our results showed that both male and female lizards had smaller territories in a habitat with greater prey biomass than lizards in habitats with less available prey, but the rates of aggressive behaviors used to defend territories did not differ among these habitats. Further, we did not find a relationship between body size and territory size, and the sexes did not differ in their relationships between food availability and territory size or behavioral defense. Together, these results suggest that differences in food availability influenced male and female territorial strategies similarly, and that territory size may be more strongly associated with variation in food resources than social display behavior. Thus, anole investment in the behavioral defense of a territory may not vary with territory quality.  相似文献   

9.
Male territorial defence is a component of many vertebrate mating systems and is often regarded as a tactic for acquiring mates. Traditionally considered within the context of overt site‐specific defence, territoriality actually may have several components which encompass a variety of behavioural tactics (e.g. post‐copulatory mate‐guarding, defence of resources that females need, defence of area around females) that underlie a mating system. The purpose of our study was to evaluate such influences on the territorial behaviour of male Columbian ground squirrels in southwestern Alberta, Canada. Males were dominant and territorial if they defended a minimum convex polygon activity range by chasing other males more within the activity range than they were chased. Subordinate males had no territory and were chased throughout their ranges, but they competed for mates by increasing chases in their activity range when nearby females were oestrous. Dominant males exhibited conditional breeding tactics, tending to chase other dominant males from their territory when nearby females were oestrous, but travelling outside their activity ranges to chase subordinate males when females were not oestrous. Although females mated first with a dominant male on whose territory they resided (and in order from oldest to youngest if several territories overlapped), mating pairs were not exclusive, as females usually mated with additional males. Males also guarded females after copulation and defended females directly just before oestrus, rather than defending territory per se during those times. Thus, males possess a repertoire of behaviours that complement site‐specific territoriality, and territory ownership serves to facilitate a first mating with females that live on the territory.  相似文献   

10.
The territorial defense hypothesis and the ecology of insular vertebrates   总被引:2,自引:0,他引:2  
Insular lizards, birds, and mammals in high-density populations often exhibit reduced situation-specific aggression toward conspecifics. This aggressive behavior can be expressed in the form of (1) reduced territory sizes, (2) increased territory overlap with neighbors, (3) acceptance of subordinates on the territory, (4) reduced aggressiveness to certain classes of conspecifics, or (5) abandonment of territorial defense. These behavioral traits can be explained by two nonexclusive hypotheses. The resource hypothesis suggests that territorial behavior is primarily adjusted to resource densities, and that resources are more abundant on islands than on the mainland (e.g., because of a lack of competing species). The defense hypothesis suggests that, in addition to any effects of resources, the costs of defense against both territorial intruders and contenders for vacant territories are higher on islands. Recent theoretical and empirical studies indicate that these behavioral changes can occur as a result of elevated defense costs, independent of resource densities. Reduced predation, more benign climates, and an absence of habitat sinks on islands would all tend to increase the density of potential intruders and contenders, and hence the costs of defense for owners of insular territories. The two hypotheses differ in their predictions about the rates of biomass production (growth or reproduction) for holders of insular territories. Reproductive and growth data from insular-mainland pairs indicate the importance of elevated defense costs, and also suggest that many insular vertebrates reallocate their breeding resources so as to produce young that are more competitive. The suite of ecological and behavioral traits exhibited by insular territorial vertebrates can best be explained by three factors operating in concert: higher available resource densities, higher defense costs, and (sometimes) a reallocation of resources to produce young that are more competitive.  相似文献   

11.
The period of territorial settlement is critical for territorial species, and the initial disputes to fix the boundaries can be energetically expensive. Territorial residents may be able to reduce defensive costs during settlement by selecting territories with landmarks at the sites of potential boundaries. We examined the effects of landmarks on defensive costs in a laboratory study of a cichlid fish, the blockhead, Steatocranus casuarius. In the landmark treatment, we placed a row of flat rocks across the centre of the aquaria; trials in the control treatment were identical but lacked landmarks. When landmarks were present, blockheads spent significantly less time in territorial defence, as they had fewer and shorter aggressive interactions with their neighbours. In addition, fights in landmark trials tended to be of lower intensity than fights in control trials: most fights in landmark trials included only low-level displays but most fights in control trials included physical contact. Both of these measures thus indicated that defensive costs were lowered by landmarks. In addition, in landmark trials typically both pairs of fish successfully established territories; in contrast, in control trials generally only one pair was able to establish a territory, with the other pair being evicted. The presence of landmarks appeared to make possible the division of the area available for settlement, with pairs establishing smaller territories than when there were no landmarks. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.   相似文献   

12.
Large male fiddler crabs sometimes help smaller neighbours to defend their territories against intruders. These coalitions occur when the helper is likely to defeat the intruder (helper larger than intruder) and the neighbour is likely to lose his territory without help (intruder larger than neighbour). Previous studies of coalitions have excluded males with regenerated claws. Such claws are weaker weapons that make the bearer competitively inferior. Here, we show that male Uca annulipes with regenerated claws are as likely as males with original claws to help their neighbours in territory defence, even though, as weaker males they potentially pay greater costs, being more likely to lose their undefended burrow. We suggest that males with regenerated claws gain greater benefits from retaining a current, small neighbour and that, as in non‐coalition fights, the regenerated claw acts as a visual bluff in the early stages of combat. Furthermore, we show that intruders with regenerated or original claws are equally likely to be attacked by a ‘helping’ neighbour. This bolsters the argument that males cannot visually differentiate between original and regenerated claws.  相似文献   

13.
Territorial behaviour in female small mammals has been proposed as a mechanism to defend limited ecological resources or their pups against conspecific infanticidal or predators. Female territorial behaviour very often is associated with reproductive activity due to the fact that frequency and intensity of aggression are exhibited mainly when females are pregnant or lactating. In vole and mice species, female territoriality would be a counterstrategy to prevent the killing of their pups by conspecific breeding females. To study whether female territoriality is a strategy for pups or nest defence against infanticidal breeding females, and whether time invested in nursing young affects aggressive response of mothers, we used the Pampean grassland mouse (Akodon azarae) as an ecological model species. We conducted resident–intruder tests between lactating females. Differences in residency time (48 vs. 72 h) of focal females in their home territory were also included in the analysis. In all cases, the pups of both resident and intruder mothers were placed with the nesting material from their reproductive cages. Resident mothers were always more aggressive than intruders and they were even more aggressive when they spent more time nursing their pups. Contrarily, intruder females exhibited the greatest values of submissive behaviours. Our results show that female territoriality of A. azarae would represent a strategy to protect pups from potentially infanticidal females. We discuss the extent of female intrasexual territoriality and its potential adaptive significance in relation to strategies which lead to increase their reproductive success.  相似文献   

14.
Male Polistes canadensis and P. carnifex aggregate along crests of prominent ridges in Santa Rosa National Park, Costa Rica. At these sites males of both species defend territories (trees and shrubs) by chasing conspecific rivals. Territories do not contain nests or resources that are collected by females. Chasing by territorial males reduces the amount of time spent by intruders in a territory. I describe and contrast male territorial behavior of both species. Some male P. canadensis are territorial while others in the same area exhibit patrolling behavior, flying from one occupied territory to another. Males of P. carnifex exhibit territoriality only. Patrolling in P. canadensis is an outcome of relatively high male density along the ridge, rendering territories in short supply, as shown by the observation that experimentally vacated territories are seized rapidly by formerly patrolling males. Due to a high intraspecific intrusion rate, territorial male P. canadensis spend less time perching and more time flying and chasing intruders from their territories than do male P. carnifex. Males of these two species also differ in the placement of their territories along the ridgeline; P. canadensis occupy territories in saddles while P. carnifex occupy those at peaktops. I show that this divergent spatial pattern is not maintained by competitive exclusion of either species by the other, and I discuss alternative explanations for their separate spatial distributions. Comparative data suggest that males are territorial because females restrict matings to within territories, and I discuss alternative hypotheses to explain this bias in female behavior.  相似文献   

15.
The standard length, diurnal activities, territory sizes, and areas over which individuals foraged and the nutritional condition of territorial and non-territorial Cynotilapia afra and Pseudotropheus zebra were compared. Results show that territorial tenureship in these fishes does not depend on the male size, implying that aggressiveness, experience and motivation are more important in the maintenance of territory. However, terri-toriality in C. afra (t = 10.93, P < 0.05) and P. zebra (t = 3.31, P < 0.05) manifests itself in the reduction of nutritional condition because territorial males engage in energetically more demanding activities, e.g. courtship displays, fertilizing ripe females and chasing intruders that trespass into their territories. Territorial C. afra (t = 4.77; P < 0.05) and P. zebra (t = 5.89; P < 0.05) also fed over significantly smaller areas and spent significantly less time feeding than did non-territorial males. The biological significance of territoriality in these fish species is therefore not food intake, but mate attraction and reproductive success for which they trade off their nutritional condition. However, there might be a nutritional threshold below which the cost out-weighs the benefit. Hence, territorial males in poorer health abandon their territories in order to regain their condition.  相似文献   

16.
Avoiding predators may conflict with territorial defence because a hiding territorial resident is unable to monitor its territory or defend it from conspecific intrusions. With persistent intruders, the presence of an intruder in the near past can indicate an increased probability of future intrusions. Therefore, following a conspecific-intrusion, territorial residents should minimize costs from future intrusions at the cost of higher predation risks. I conducted experiments with males of the territorial lizard Tropidurus hispidus recording approach distance (distance between predator and prey when the prey escapes) and time to re-emergence from a refuge after hiding. Past aggressive interactions affected anti-predator behaviour: lizards re-emerged sooner (compared to a control) when the predator attacked 5 min after an aggressive encounter. If the predator attacked while an aggressive encounter was ongoing, there was also a reduction in approach distance. The results are consistent with an economic hypothesis which predicts that T. hispidus incur greater predation risks to minimize future territorial intrusion; additionally they show that the effects of past and ongoing aggressive interactions are different, consistent with the minimization of present intrusion costs. These results are relevant for studies of the changes in aggressive behaviour due to changes in the social environment and for studies of the costs and (co) evolution of aggressive and anti-predator strategies.  相似文献   

17.
Differential responses to neighbours and strangers (the dear enemy phenomenon) and individual recognition presumably evolve to reduce costs of territorial defence. Territorial residents have been found to demonstrate reduced aggression towards neighbours wherever they are encountered along that resident's territory boundary except for when the neighbour is displaced to the boundary opposite the shared boundary. In this new location, the displaced neighbour represents a greater threat to the resident's territory ownership, and should be treated as equally aggressive as a stranger. Finding increased aggression towards displaced neighbours has been interpreted as individual recognition, but these results do not provide sufficient evidence to rule out the possibility that the resident sees the neighbour out of its normal context as just another stranger. We tested the hypothesis that territorial collard lizards can individually recognize neighbours and will increase aggression towards them as the threat to territorial ownership increases. Resident males treated neighbours that had been moved to the opposite boundary as equally aggressive as strangers. However, residents responded more aggressively towards strangers than towards neighbours on natural territories (the dear enemy phenomenon) and also in neutral arena encounters. Our results suggest that resident male collared lizards are able to recognize individuals regardless of context and respond to them according to the threat that they pose. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.   相似文献   

18.
Aggression serves a great variety of social functions, one of which is protection of individual territories from intruders. Territorial males of many antelope species show aggressive noncontact displays, and only rarely fight. It has been suggested that ungulate males tend to have more frequent and longer aggressive interactions with rivals of similar age or social status than with males of dissimilar status. In the present paper, we test whether territorial and non-territorial males behave in a similar manner and avoid fights, and whether or not they preferentially direct aggressive and longer agonistic interactions towards males of similar age or social status, rather than towards other classes of males. We found that territorial males usually avoided straight fights with peers, and instead mainly used noncontact displays in aggressive interactions. In contrast, non-territorial males used fights considerably more often, especially during the onset of territoriality in April to May, when these males had their most frequent aggressive interactions. Territorial bucks aggressively interacted most frequently with non-territorial males and significantly less often with other territorial males, but agonistic noncontact displays between territorial males lasted the longest. In contrast, non-territorial males addressed their aggressive noncontact displays and fights most often to peers and less frequently to sub-adults. Asymmetry in the social status of territorial vs. non-territorial males was likely responsible for the distinctively different agonistic behaviors shown by the two types of males, which in turn are likely due to the different costs and benefits each male can accrue from these aggressive interactions.  相似文献   

19.
Territorial songbirds often match the song features or singing patterns of rivals, commonly as an aggressive signal. Most studies of song matching have been on Northern Hemisphere species with short lifespans and high song rates, but vocal matching is predicted to be affected both by longevity and territorial stability. We studied song matching in males of the white-browed scrubwren, Sericornis frontalis, a long-lived, sedentary, territorial Australian songbird. We quantified natural song rate and diversity, and then conducted three playback experiments to test: (a) whether males match by song type; (b) how they respond physically and vocally to territorial intrusion; and (c) whether they match by song length, and use it as an agonistic signal. Males naturally had very low song rates, singing on average less than three times per hour, and moderate repertoires, with an estimated mean of 17.5 song types for individual males. Males did not engage in extended counter-singing bouts. The first experiment showed that males matched the song type of immediate neighbours almost 90% of the time, if that type was in their repertoire. The remaining experiments revealed that song-type matching was an aggressive signal; males responded more aggressively to, and were more likely to match, playback simulating a neighbour's territorial intrusion than song from their shared boundary. Males did not match songs by length, but they produced longer songs after simulated intrusion. Males also responded more aggressively to playback of longer songs that simulated intrusion, but less aggressively to longer songs from the territory boundary. Overall, we show that sedentary, long-lived songbirds with low song rates, can use song-type matching as an aggressive signal to communicate with neighbours and intruders. Song length had a different role in communication, possibly related to individual quality or territory ownership.  相似文献   

20.
Many animals defend territories against conspecific individuals using acoustic signals. In birds, male vocalizations are known to play a critical role in territory defence. Territorial acoustic signals in females have been poorly studied, perhaps because female song is uncommon in north‐temperate ecosystems. In this study, we compare male vs. female territorial singing behaviour in Neotropical rufous‐and‐white wrens Thryothorus rufalbus, a species where both sexes produce solo songs and often coordinate their songs in vocal duets. We recorded free‐living birds in Costa Rica using an eight‐microphone Acoustic Location System capable of passively triangulating the position of animals based on their vocalizations. We recorded 17 pairs of birds for 2–4 consecutive mornings and calculated the territory of each individual as a 95% fixed kernel estimate around their song posts. We compared territories calculated around male vs. female song posts, including separate analyses of solo vs. duet song posts. These spatial analyses of singing behaviour reveal that males and females use similarly sized territories with more than 60% overlap between breeding partners. Territories calculated based on solo vs. duet song posts were of similar size and similar degrees of overlap. Solos and duets were performed at similar distances from the nest for both sexes. Overall, male and female rufous‐and‐white wrens exhibit very similar spatial territorial singing behaviour, demonstrating congruent patterns of male and female territoriality.  相似文献   

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