Territorial behavior of the Korean wood-feeding cockroach, <Emphasis Type="Italic">Cryptocercus kyebangensis</Emphasis>

We investigated the territorial behavior of Cryptocercus kyebangensis from the viewpoint of offspring protection. Aggressive behavior against adult intruders occurred more frequently in resident adults with their young than those without their young. Resident adults without offspring more frequently attacked intruders of the same sex than those of the opposite sex, whereas those with offspring attacked regardless of the sex of adult intruders. In paired residents, cooperative territorial behavior occurred in a higher proportion in pairs with offspring than those without offspring. Responses of residents to nymphal intruders depended on the head capsule size of the intruders. Resident adults rarely attacked intruders smaller than their own offspring but frequently attacked larger intruders. Resident nymphs frequently attacked intruders smaller than themselves, whereas they escaped or showed jerking behavior when they encountered larger intruders. Electronic Publication     

Effects of resource distribution on activity and territory defence in house mice, Mus domesticus

How much an animal invests in defending a territory depends, in part, on the quality, quantity and distribution of resources, but do animals target their investment in defence within the territory according to the location of different resources? We examined whether wild-caught male and female house mice increased aggression towards intruders of the same or opposite sex when in areas containing their food/water or nest site. Pairs of resident mice were established in enclosures consisting of two areas. In split enclosures, food and water were in one area and a nest site in the other, while in clumped enclosures all resources were in one area with no resources in the other. Residents of both sexes were quicker to attack intruders, and were more aggressive, in areas containing resources than in an area containing no resources. There was no difference in aggression between areas containing food/water, a nest site or both. The effect of resources on territorial defence was due largely to changes in aggression against intruders of the opposite sex. Residents, especially males, attacked intruders of their own sex more than those of the opposite sex. This bias was much stronger when no resources were present, owing to low aggression against opposite-sex intruders in the absence of resources. There were no differences in intruder aggression, and the relative size of the intruder (heavier or lighter than the resident) had no effect on resident aggression. Residents also spent less time out of their nest in clumped enclosures owing to low activity in the nonresource area.     

Territorial behaviour reflects sexual status in groups of false clown anemonefish (Amphiprion ocellaris) under laboratory conditions

Anemonefishes (genus Amphiprion) are well known as protandrous sex-changing fish. They live symbiotically with sea anemones and form a social unit that consists of a breeding pair and several sexually immature individuals with ambisexual gonads. In general, the hierarchy of the social rank is strictly maintained in a group, and this hierarchy is considered to be one of the critical factors that induce sex change or sex differentiation in anemonefishes. Field observations have shown that behavioural characteristics of anemonefishes vary depending on the environment. In this study we examined the differences in territorial aggression directed at conspecific intruders according to sexual status in the false clown anemonefish, Amphiprion ocellaris, under laboratory conditions. Resident–intruder tests were conducted on groups consisting of three anemonefish: a dominant female, a second-ranked male and a subordinate ambisexual individual. Females and males showed more aggressive behaviour than did ambisexual individuals, and behaviours were specifically directed at intruders of the same sexual status, not at those of the opposite sex. Notably, the female residents displayed fierce aggressive behaviours particularly toward female intruders, whereas male residents engaged more often in side-by-side swimming with male intruders than with female or ambisexual intruders. Ambisexual intruders were threatened by all residents, rarely attacked. We observed only few aggressive acts towards tank mates. These results suggest that sexually mature resident anemonefish in a confined space, such as under laboratory conditions, perceive intruders of the same sexual status as competitors for reproductive status and thus, dominant social rank.     

Responses of a plethodontid salamander to conspecific and congeneric intruders

After inhabiting their home sites for a few days, both males and females of the salamander Desmognathus monticola directed agonistic behaviour toward intruding salamanders. Resident D. monticola were significantly more aggressive toward D. fuscus intruders than toward conspecific ones, although both conspecific and congeneric intruders were frequently attacked. Responses of D. monticola residents were independent of sex, and no difference was found in the behaviour of residents toward conspecific male and female intruders. Small D. fuscus intruders were attacked and injured more frequently than similar-sized D. monticola intruders and, in general, smaller-sized salamanders were the victims of aggressive attacks more frequently than larger ones. The proportions of bites on the head and tail of intruders were similar, although injuries were much more frequent to the tail than to the head. This species-related and size-related agonistic behaviour may be functional in maintaining the spatial distribution of species in communities of Desmognathus salamanders.     

Maternal aggression as a deterrent to infanticide in Peromyscus leucopus and P. maniculatus

Adult male and female white-footed mice (Peromyscus leucopus) and deermice (P. maniculatus) from sympatric populations commit infanticide on neonates of either species. A laboratory experiment was conducted to determine whether aggression in maternal females could effectively deter infanticidal intruders. Females of both species exhibited post-partum aggression and dominated intruders of either sex or species in 83 of 88 trials. In the absence of the mother, neonates were attacked by intruders in 82 out of 84 trials. Maternal females were considerably more aggressive than anoestrous females. The implications for female territorially as a mechanism to prevent infanticide are discussed.     

Intruders in Nests of the Spotless Starling: Prospecting for Public Information or for Immediate Nesting Resources?

The prospective behaviour for nests by conspecific intruders may be a strategy to gather public information for future reproduction or to secure resources for immediate reproduction. Although the relationship between the sex, age and breeding experience of intruders and the sex and age of residents may be crucial for understanding the significance of nest‐prospecting behaviour, a precise determination of these traits has rarely been addressed in field studies. In a free‐ranging population of spotless starlings ( Sturnus unicolor), inexperienced birds were the predominant intruders. The ratio of male‐to‐female intruders was higher among birds without previous breeding experience, although our results did not allow us to determine whether more inexperienced males or females intruded nests. The average annual rate of intrusion of either sex was strongly correlated with the number of fledglings produced in the study colony the previous year, which seems to indicate that the inexperienced intruders were, in general, yearlings. The proportion of the sexes among the intruders in each study year was correlated with the average age of male owners but not with the age of female owners. This result suggests that the presence of intruding males in the nests was affected by an increased opportunity to find vacancies in nesting territories as resident males became older. This finding supports the hypothesis that, in most cases, birds visited nests to obtain personal information about nesting resources. However, the intrusion rates were higher when there were nestlings than during the incubation period, which suggests that collecting public information was also involved in the intruding behaviour. Both sexes regularly attacked intruders, which implies that the intruders inflicted some kind of cost on the owners. Female floaters were more frequently chased, probably because they regularly lay parasitically in conspecific nests.     

Sexual dimorphism in responses to unfamiliar intruders in the tamarin, Saguinus oedipus

Aspects of social structure in group-living species can be inferred by observing the responses of individuals to unfamiliar animals. This study examined the responses of mated pairs of cotton-top tamarins (Saguinus oedipus oedipus) to presentation of either unfamiliar conspecifics or members of a related tamarin species, the saddle-back tamarin (Saguinus fuscicollis fuscicollis). Male and female responses to intruders differed: resident males threatened, displayed piloerection, approached, and attacked intruders, especially males, while resident females showed increases in suprapubic scentmarking in the presence of intruders. Both males and females discriminated between the species of intruders, exhibiting more threats, scent-marking, piloerection, and approaches in the presence of conspecific intruders. There are pronounced sex differences in the signals and behaviour patterns that are elicited in an aggressive or territorial encounter with unfamiliar conspecifics in this monogamous primate.     

Colony defense in Damaraland mole-rats, Cryptomys damarensis

I examined defense against conspecific intruders in coloniesof the Damaraland mole-rat. I introduced foreign animals tocaptive colonies and examined the effect of the sex and breedingstatus of the intruder on the sex and breeding status of thosethat defended. Intruding mole-rats were almost always attacked.Colony defense against intruders was carried out almost exclusivelyby dominant, reproductive individuals and was directed primarily toward individuals of the same sex. Nonbreeding, subordinatemole-rats participated only rarely. These results contrastsharply with colony defense in the related naked mole-rat andsuggest that investment in defense activities in Damaralandmole-rats primarily reflects repulsion of potential competitorsfor breeding status. Subordinate mole-rats may lack incentiveto participate in potentially costly defense, as they are unlikelyto gain reproductive opportunities within the colony and probablygain fitness primarily through dispersing to breed.     

Acquisition of food by fledgling Egyptian Vultures Neophron percnopterus by nest-switching and acceptance by foster adults

In a dense population of Egyptian Vultures Neophron percnopterus in northern Spain, nest-switching occurred between nearby nests. Fledglings obtained more feeds by intruding and pirating than by remaining in their own nests during the same period. Intruder fledglings showed the same behaviour with Foster adults attacked intruder fledglings significantly less than they attacked alien adults and immatures but more frequently than they attacked their own fledglings. Effective rejection behaviour by foster adults to intruders may not have evolved as Egyptian Vultures usually nest at lower densities than that found in the study area.     

Postpartum aggression in mice: Experiential and environmental factors

Six experiments were conducted to assess the influence of duration of lactation, the presence of young, and the stimulus characteristics of intruder animals upon postpartum aggression of mice. The first experiment showed that postpartum aggression toward conspecifics was highest between Day 3 and Day 8, declined between Day 9 and Day 14, and was present toward males but absent toward females between Day 15 and Day 21 of the lactation period. Experiment 2 showed that lactating mice rarely attacked conspecifics to which they had been previously exposed but would readily attack strangers. Experiment 3 and 4 demonstrated that lactating animals never attacked intruders when tested 5 hr after pup removal. However, placement of young behind a wire partition in the home-cage for 5 hr or replacement of the offspring for as little as 5 min following 5 hr of separation restored postpartum aggression. The fifth experiment showed that 1- and 10-day old intruders were seldom attacked while intense aggression was directed against 14- and 20-day old intruders. Finally, Experiment 6 demonstrated that 14-day old intruders whose hair was removed were rarely attacked.     

The Role of Wing Coloration in Sex Recognition and Competitor Recognition in Rubyspot Damselflies (Hetaerina spp.)

The decision rules that animals use for distinguishing between conspecifics of different age and sex classes are relevant for understanding how closely related species interact in sympatry. In rubyspot damselflies (Hetaerina spp.), the red wing coloration of mature males is hypothesized to be a key trait for sex recognition and competitor recognition within species and the proximate trigger for interspecific male–male aggression. We tested this hypothesis by manipulating the wing coloration of tethered conspecific intruders and measuring the responses of territory holders of three species in the field. As predicted, covering the red spots of mature males with black ink nearly eliminated territorial responses, and in some cases, territorial holders clasped the blackened males as if they were females. Adding red spots to female wings triggered territorial responses and nearly eliminated sexual responses. Immature males with artificial red spots were attacked at the same rate as mature male intruders, and much more frequently than were immature male controls. The results varied somewhat by species. In H. titia, the only species of Hetaerina with substantial black wing pigmentation, the effects of blackening the red spots of intruders varied both geographically and seasonally. But even when blackening the red spots of male intruders did not reduce the aggressive response of H. titia territory holders, adding artificial red spots to female wings elicited aggressive responses and nearly eliminated sexual responses. The results of this study further strengthen the evidence that interspecific aggression in Hetaerina results from overlap in territorial signals and that the derived black wing pigmentation of H. titia reduces interspecific aggression.     

Risk-sensitive resource defense in a territorial reef fish

As coral reefs are home to dense aggregations of a variety of species, aggressive territoriality is often a critical component of individual behavior. Identification and assessment of the risk posed by intruders is crucial to defending a territory, and fishes on coral reefs have been found to attend to body shape, body size, and coloration when responding to intruders. We examined the extent to which dusky damselfish (Stegastes adustus) discriminate among distinct categories of intruders by measuring the distance at which a fish attacks an intruder and the relative intensity and frequency of those attacks. We found that S. adustus discriminated among perceived threats, attacking conspecifics more intensely and more often than egg-predators and herbivores, and showing a trend of attacking those groups more often than invertebrate-feeders, which do not compete with damselfish for resources. Furthermore, territory holders attacked initial-phase wrasses from a farther distance than terminal-phase wrasses, suggesting that they can discriminate among classes of individuals within a species other than their own. Dusky damselfish thus exhibit the ability to make fine distinctions among intruders in a diverse ecosystem.     

Depth-related differences in territory size and defense in the herbivorous cichlid,neolamprologus moorii, in lake tanganyika

Neolamprologus moorii, a small herbivorous cichlid of Lake Tanganyika, defends its territory against food competitors. Territory size and territorial defense were compared between two sites at different depths. In the shallow site, more fishes intruded into the territories than in the deep site, owing to the higher density fish population in the former. The territory holders in the shallow site attacked intruders more often, although the ratio of attacks to number of intruders was similar to that of the deep site. The territories in the deep site were larger and contained a greater amount of algae than the shallow site. The larger territory size in the deep site may have resulted from the lower cost of territory defense due to the lower competitor density and from the need to maintain a greater amount of food resources to compensate for lower algal productivity.     

Effects of residence,aggressive experience and intruder familiarity on attack shown by male mice

Two groups of 12 mature male Swiss Morini strain mice were matched for aggressiveness on the basis of their response to anosmic docile male intruders after 24 h individual housing in a large defensible cage. One group was subsequently individually housed without disturbance for a further 12 days whereas the second group was exposed to a new anosmic intruder every 3 days over this same period. All mice were subsequently retested against anosmic intruders for a 10 min period. Animals which had had the opportunity to fight repeatedly showed more attack than individuals lacking such experience. A second experiment contrasted the responses of dominant isolates to the same or a different anosmic intruder, 10 min after a successful attack. Familiar intruders were attacked less vigorously than unfamiliar mice. It seems likely that the defeated mouse becomes less potent as a stimulus eliciting attack as the resident becomes habituated to it. Conversely, unfamiliar intruders evoke aggressive reactions by residents despite the prolonged fights they had had before final testing. These experiments provide little support for the appetence view of aggression in mice (i.e. the view that aggressiveness is augmented by depriving the animal of the opportunity to fight) or the view that this phenomenon is simply a consequence of “social deprivation”.     

Riverside wren pairs jointly defend their territories against simulated intruders

Duets are a jointly produced signal where two or more individuals coordinate their vocalizations by overlapping or alternating their songs. Duets are used in a wide array of contexts within partnerships, ranging from territory defence to pair bond maintenance. It has been proposed that pairs that coordinate their songs might also better coordinate other activities, including nest building, parental care and defending shared resources. Here, we tested in the riverside wren (Cantorchilus semibadius), a neotropical duetting species that produces highly coordinated duet songs, whether males and females show similar responses to playback. During territorial disputes in songbird species, individuals tend to direct their attention towards same‐sex territorial intruders, but this bias might be less pronounced in duetting species. We performed a dual‐speaker playback experiment to examine how mated individuals respond to speakers broadcasting female‐versus‐male duet contributions. We found that riverside wrens have high levels of converging behaviour by duetting and remaining in close proximity to one another when responding to simulated paired intruders. Males and females spent more than 80% of their time less than 1 m apart while defending their territory. Both individuals in a pair aggressively engaged with both male and female simulated trespassers by approaching equally close and spending equal time near the two speakers. These results suggest that both sexes perceive a paired territorial intrusion as a similar threat and that both partners are highly invested in defending the shared resources. This study is one of the few to demonstrate equal attention and aggression from mated pairs towards simulated same‐sex and opposite‐sex intruders. We suggest that pairs responding together, in close proximity to one another, might be favourable in duetting species when defending the territory because maintaining a close distance between partners facilitates the extreme coordination of their joint territorial signals.     

Sex-specific territorial responses in Tawny Owls Strix aluco

A territorial intruder often poses more of a threat to the territory holder of the same sex as itself. As territorial aggression is associated with costs, aggression shown by birds towards territorial intruders of the opposite sex deserves investigation. This behaviour could arise due to the reproductive value of a mate or through mutualism between members of a pair. We investigated these hypotheses by presenting mated pairs of Tawny Owls Strix aluco with playback of male calls, female calls and a male and female duetting, and recording the number and intensity of responses by the male and female territory holders. Females responded significantly more often to female than to male calls. Males responded equally often to male and female playback. Males which had previously bred successfully with their mate were significantly more likely to respond to female playback in the spring, which suggested males were responding to female playback due to the reproductive value of their mate. There was no evidence of mutualism between members of a pair.     

The effect of the type of opponent in tests of murine aggression

The interactions between isolated resident mice and anosmic, juvenile, submissive and dominant intruders were studied; the responses of intruder mice to being attacked and the distribution of bites on these subjects were investigated. Anosmic, juvenile and (trained) submissive intruders did not attack residents; they did, however, show differences both in thir responses to being attacked and their ability to avoid bites to particular regions of their body surface. These differences are likely to make it difficult to extrapolate between tests that employ different types of intruder. Trained dominant intruders showed substantial attack on resident mice and a very different pattern of attack was generated by this type of encounter. The study confirms that, while “standard opponents” are useful means of assessing aggressive motivation, the pattern of fighting they produce differs depending on their background.     

Aggressive behaviour in Montagu’s harrier Circus pygargus during the courtship period

Aggressive behaviour of Montagu’s harrier was observed during the pre-laying period in the 1992–1995 seasons on the calcareous marshes of Chelm in Eastern Poland. In total, 435 flights performed by 24 pairs of individually marked harriers were analysed. All flights were performed in relation to the territory of defence. Males performed 61% of aggressive interactions while females performed 39%. Intraspecific and interspecific aggression comprising direct attacks (58%), escorts (13%), pursuits (28%) and “mirror behaviour” (1%) were described for the first time. Interspecific aggressive behaviour occurred only near territory boundaries. The main intruders were marsh harriers Circus aeruginosus breeding on the same marshes. Occasionally, harriers attacked curlews Numenius arquata, short-eared owls Asio flammeus, magpies Pica pica, hooded crows Corvus corone cornix, common buzzards Buteo buteo or hobbies Falco subbuteo. Four cases of communal mobbing were observed. Three to five males from the neighbourhood attacked the intruders together. Intraspecific aggressive behaviour was observed in the pair territory, near the boundary or at a distance of up to 100 m from the defended area. Aggressive interactions performed by both sexes were more frequent towards birds of the same sex. Females defended their territories more aggressively against females. Similarly, males were more aggressive against males. All cases of aggressive behaviour were observed near harrier territories. Aggressive relations between birds outside breeding territories or when foraging around marshes were not observed.     

A high aggression strategy for smaller males

Male-male conflict is common among animals, but questions remain as to when, how and by whom aggression should be initiated. Factors that affect agonistic strategies include residency, the value of the contested resource and the fighting ability of the two contestants. We quantified initiation of aggression in a fish, the desert goby, Chlamydogobius eremius, by exposing nest-holding males to a male intruder. The perceived value of the resource (the nest) was manipulated by exposing half of the residents to sexually receptive females for two days before the trial. Resident male aggression, however, was unaffected by perceived mating opportunities. It was also unaffected by the absolute and relative size of the intruder. Instead resident aggression was negatively related to resident male size. In particular, smaller residents attacked sooner and with greater intensity compared to larger residents. These results suggest that resident desert goby males used set, rather than conditional, strategies for initiating aggression. If intruders are more likely to flee than retaliate, small males may benefit from attacking intruders before these have had an opportunity to assess the resident and/or the resource.     

The influence of target sex and dominance on evoked attack in Rhesus monkeys

Attack behaviors were elicited by the electrical stimulation of the hypothalamus of freely moving male rhesus monkeys. The influence of the sex and dominance status of target animals on the evoked response was measured. Animals offered as targets were dominant males and females and subordinate males and and females. Males were attacked more often than females and subordinate targets of either sex were attacked more often than dominant targets. A comparison was made between spontaneous and evoked attack behavior.