The influence of predation risk on threat display in great tits

In wintering birds, conflicts over food are often resolved bythreat displays. For displays to be effective, there oughtto be a cost associated with displaying. We investigated whetherincreased vulnerability to predators due to reduced vigilancecould be such a cost. Conflicts ought then to be resolved usingfewer or less intense displays in conditions of high risk. We also looked for differences between dominants and subordinatesin their reaction to risk. Because there is considerable evidencethat subordinate wintering birds forage in riskier places thandominants, one might expect dominants to be less successfulin conflicts under high predation risk. In our experiment,nine flocks of four or five wintering male great tits were keptin outdoor aviaries. In the predation risk treatment, a stuffedpygmy owl was briefly shown before birds were allowed accessto a feeder. In the control treatment the owl did not appear.The predator presentation caused a reduction in the amountof aggression shown by subordinates, whereas for dominants there was no statistically significant change. Dominants were at leastas successful in subduing subordinates under high risk as underlow risk. A possible interpretation is that our experimentreflected a natural foraging situation for great tits, whereephemeral resources can appear unpredictably. In such situations,dominants may need to be bold to gain priority of access even under increased risk of predation, whereas a subordinate wouldgain little by risking a conflict with small chances of winning.     

Multiple Cues in Status Signalling: The Role of Wingbars in Aggressive Interactions of Male House Sparrows

During aggressive interactions, animals may signal their competitive ability by various ornaments referred to as badges of status. The use of a single badge predicting dominance rank occurs in many vertebrate species. However, animals often display multiple ornaments that may convey information about either different or the same aspects of the signaller's quality, or alternatively, may serve as signal amplifiers. We observed the fighting behaviour of male house sparrows in two captive flocks to investigate whether they may use multiple cues in status signalling during aggressive interactions. Beside the status‐signalling bib, male sparrows possess a conspicuous white wingbar that they often display upon aggressive encounters. We tested whether bib size and the wingbar's conspicuousness (i.e. its achromatic contrast with the neighbouring dark feathers) or its area predicted success in various aspects of fighting. We found that bib size strongly predicted overall fighting success (i.e. proportion of fights won) and defence success (i.e. proportion of successful defences out of all attacks received). Wingbar conspicuousness was positively related to defence success after controlling for the effect of bib size in multivariate analyses. Furthermore, displaying the wings also tended to improve the birds’ success in defence but not in attack. Wingbar area was unrelated to any measured aspect of fighting ability. We suggest that bib size and wingbar conspicuousness may convey multiple messages on fighting abilities, specifically on overall aggressiveness and defending potential, respectively. Alternatively, wingbars may serve as amplifiers for the wing displays of aggressive motivation. Thus, male sparrows may use multiple cues in assessing the competitive ability of opponents during social interactions.     

Aggression in shorebirds in relation to flock density and composition

The aggressive interactions of Turnstones and Purple Sandpipers feeding in wintering mixed-species wader flocks could be classified into those involving food and those involving space. All observed interspecific encounters were of short duration and were initiated and won by the larger species; the majority did not involve food and were resolved by low-intensity displays. Intraspecific interactions (a greater proportion of which involved food) were also resolved quickly and were usually won by the initiator. Space-related encounters between conspecifics were more likely to be resolved than food-related encounters just by threat displays. Aggression rates increased with flock densities. However, the increase in aggression with density was dependent on the species composition of the flock: both Turnstones and Purple Sandpipers were more likely to be involved in fights (both over food and over space) with conspecifics than with other species, indicating that the aggression costs of flocking were less in mixed-species flocks.     

Digital video playback and visual communication in lizards

Experimental analyses of dynamic visual signals have to overcome the technical obstacle of reproducing complex motor patterns such as those found in courtship and threat displays. Video playback offers a potential solution to this problem, but it has recently been criticized because of sensory differences between humans and nonhuman animals, which suggest that video stimuli might be perceived as deficient relative to live conspecifics. Quantitative comparisons are therefore necessary to determine whether video sequences reliably evoke natural responses. Male Jacky dragons, Amphibolurus muricatus, compete for territories using complex displays delivered in a rapid stereotyped sequence. We evaluated video playback as a technique for studying this visual signal. Digital video sequences depicting a life-sized displaying male were indistinguishable from live male conspecifics in the rate and structure of aggressive displays evoked. Other measures of social behaviour suggested that video stimuli were more effective in this context. Lizards produced significantly more appeasement displays and had higher rates of substrate licking and locomotor activity in response to video playback than to confined male opponents, which failed to produce aggressive displays. Lizards tracked temporal changes in the display rate of video stimuli and were also sensitive to individual differences in morphology and behaviour between video exemplars. These results show that video stimuli are appropriate for the experimental analysis of Jacky dragon aggressive displays. We compare the potential shortcomings of video playback with those of other techniques and conclude that no approach offers a panacea, but that several have complementary characteristics. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

An ambiguous function of an alarm pheromone in the collective displays of the Australian meat ant,Iridomyrmex purpureus

Alarm pheromones, which have been documented in many species of ants, are thought to elicit responses related to aggressive or defensive behaviour. The volatile odour 6-methyl-5-hepten-2-one is described as an alarm pheromone in several species of ants, including the Australian meat ant, Iridomyrmex purpureus. The alarm pheromone is released by displaying workers that aggregate in the characteristic collective display grounds, located mid-way between colonies or near contested food trees. Workers are typically more aggressive at the latter location, and the alarm pheromone may regulate the collective level of aggression. We investigated this possibility by exposing displaying workers to synthesised alarm pheromone 6-methyl-5-hepten-2-one in a field experiment, and measuring their aggressive behaviour. We found no evidence that exposure to synthesised alarm pheromone caused changes in the aggressive level of workers. Subsequent field experiments revealed that the pheromone functions as an attractant, thereby increasing the density of displaying workers. More densely populated workers also display more aggressively, indicating that the interaction rate of displaying workers may determine the level of aggression in collective displays. This underlying mechanism can explain why displaying ants are more aggressive at the more densely populated food-tree locations than those displaying at locations midway between two neighbouring colony nest sites.     

Behavioural determinants of agonistic success in invasive crayfish

Ecosystems today increasingly suffer invasions by multiple invasive species, some of which may share similar advantageous life history traits and ecological niche. In such cases, direct competition can influence invasion success of both species, and provide insights into competition without co-evolution in species equally novel to the environment. We used two widespread crayfish invaders of freshwater ecosystems of Europe, signal crayfish (Pacifastacus leniusculus) and spiny cheek crayfish (Orconectes limosus), to investigate how behavioural decisions in agonistic encounters contribute to competitive advantages in the absence of adaptation to either opponents or an environment. In direct competition against novel but comparable opponents, the key factor for establishing clear dominance of P. leniusculus in interspecific bouts was its greater tendency towards continued engagement in high-intensity fights. With O. limosus individuals consistently retreating from staged bouts as fights became more intense, P. leniusculus individuals did not need to adapt their strategy to be successful, suggesting that their agonistic behaviour intrinsically predisposed them to win. While both species are detrimental to invaded ecosystems, our results indicate that aggressive behaviour of P. leniusculus against unfamiliar opponents could allow it to more easily outcompete other comparable species and consequently present a potentially greater threat for native ecosystems.     

Aggressive biases towards similarly coloured males in Lake Malawi cichlid fishes

In haplochromine cichlids, female mate choice based on male nuptial coloration has played an important role in speciation. Recent studies suggest that male coloration strongly influences the distribution of these fishes based on male-male aggression; males direct more aggression towards similarly coloured opponents while tolerating differently coloured individuals. We explored the role of male nuptial colour in aggression among the mbuna of Lake Malawi, examining aggression by male Metriaclima mbenjii, the red top cobalt zebra, towards conspecific opponents, similarly coloured heterospecific opponents and differently coloured heterospecifics. In trials in which focal males were offered a single opponent, while the total number of aggressive behaviours did not vary among opponent species, the types of behaviours did; focal males directed more lateral displays towards conspecifics than towards the other opponent species. When focal males were offered two opponents simultaneously, M. mbenjii directed more aggressive behaviours and more lateral displays towards similarly coloured opponents, regardless of species. Furthermore, when offered a conspecific and a similarly coloured opponent simultaneously, there were no differences in behaviour towards either opponent. Thus, nuptial coloration is used by males to identify competitors, and it suggests that male-male aggression may have also been an important diversifying force in speciation in rock-dwelling Lake Malawi cichlids.     

The relationship between social status and resting metabolic rate in great tits (Parus major) and pied flycatchers (Ficedula hypoleuca)

The oxygen consumption of great tits (Parus major) in winter flocks and of male pied flycatchers (Ficedula hypoleuca) in breeding condition, was measured. Resting metabolic rate was significantly correlated with dominance rank, as expressed by the width of the breast-stripe in the great tits and the darkness of the plumage on the head and back, or date of pair-formation, in males of the pied flycatcher. Individuals with the highest metabolic rates were the most dominant ones. Similarly, heart weight, relative to body weight, in dominant great tits was greater than that in lower-ranking birds.     

Interspecific aggression declines seasonally in breeding great tits Parus major

During the breeding season, great tits show aggression to protect their nest from intra‐ and interspecific intruders. Aggression is a labile trait that can be plastically expressed as a result of individual differences (e.g., personality), seasonal gradients in the costs and benefits of aggression, or other environmental components (e.g., number of competitors). Competitors may try to take over great tit nests, because the number of suitable nesting sites is limited, and great tits may guard high quality territories. Taking over a great tit nest may be especially fruitful in early phenological stages (egg laying) when great tits frequent their nests less often. However, great tits may compensate for this vulnerability by being more aggressive toward intruders during early nesting stages, a pattern that has already been established in an intraspecific context. Previous studies have shown that interspecific intruders were most likely to die from great tit aggression during great tit egg laying, suggesting great tits may also be more aggressive during this phase in an interspecific context. Here, I tested this hypothesis with simulated territorial intrusions in great tit territories using taxidermized blue tits Cyanistes caeruleus (hereafter called blue tit models). Great tit aggression (number of calls and approach distance toward blue tit model) was assayed during egg laying, incubation, and chick rearing in the breeding season of 2014. Although sample size was low due to a high fraction of non‐responders (n = 44 out of 89 assays across 26 out of 35 individuals), I found that great tits showed a seasonal decline in aggressiveness, which is congruent with intraspecific results on this study species. I discuss my findings in the context of differential adjustment to climate change between interspecific competitors.     

Ready to Fight: Reliable Predictors of Attack in a Cooperatively Breeding,Non‐Passerine Bird

Signal reliability is a major focus of animal communication research. Aggressive signals are ideal for measuring signal reliability because the signal referent – attack or no attack – can be measured unambiguously. Signals of aggressive intent occur at elevated rates in aggressive contexts, predict subsequent aggression by the signaler, and elicit appropriate responses from receivers. We tested the ‘predictive criterion’ in smooth‐billed anis, Crotophaga ani, by broadcasting one of two playback types (‘ahnee’ calls only or ‘ahnee + hoot’ calls), presenting a taxidermic mount, and observing the animals’ behavior. Based on the hypotheses that ‘hoot’ calls and ‘throat‐inflation’ displays signal aggressive intent, we predicted that they would be associated with attack, and that signaling rate would increase over the time period leading up to an attack. Indeed, both hoots and throat‐inflation displays reliably predicted attack. The second prediction, that signaling rate increases in the time leading up to attack, was strongly supported for throat‐inflation displays, which increased over the pre‐attack period in both treatments. Hoots increased over the pre‐attack period in ahnee playbacks but not in ahnee + hoot playbacks. Hierarchical signaling systems are characterized by early, less‐reliable predictors of attack, and later, more reliable predictors of attack. During both natural and simulated interactions, the more‐reliable throat‐inflation display tended to precede the less‐reliable hoot call, suggesting that this signaling system is not hierarchical. In a comparison of 11 putative signals of aggressive intent in birds, the throat‐inflation display had the second highest mutual information (reduction in uncertainty) among visual signals and non‐passerine signals while hoots had below‐average mutual information. Natural observations indicate that both hoots and throat‐inflation displays occur in the context of aggressive between‐group encounters, and hoots also occur during within‐group interactions. Throat‐inflation displays appear to be reliable indicators of aggressive intent, but the function of hoot calls is less clear.     

Reconciliation following aggression in patas monkeys,Erythrocebus patas

Following intra-group aggression, obvious conciliatory displays are absent from the behavioural repertoire of patas monkeys, Erythrocebus patas, while many other Old World primate species show special reconciliation gestures. When 10-min focal-animal samples that began after spontaneous aggression were compared with matched-control samples, captive adult female patas followed up on aggressive interactions, interacting sooner and more often with former opponents during post-conflict observations than during matched-control observations. Almost one-third of post-conflict observations included affiliative behaviour between former opponents, which is termed reconciliation. Matrilineally related opponents were more likely to reconcile with one another than were unrelated animals. No effect of the dominance hierarchy on tendency to reconcile was found. Thus, patas monkeys showed general patterns in post-conflict behaviour that were similar to those seen in other primates previously investigated.     

"Conventional" Signals in Avian Agonistic Displays: Integrating Theory, Data and Different Levels of Analysis

We present an integration of communication theory and data, drawing on examples from titmice (Aves: Paridae). We suggest how display actions such as lifting the head, raising the nape feathers, crest erecting and spreading the wings, act in agonistic communication when physical contact between opponents is rare. We propose that such displays largely act as strategic choice handicap signals. By giving these displays the signaller is believed to incur costs which underwrite the reliability of the signals; it may strategically increase these costs (for example by display repetition or adding additional elements) to signal its condition, motivation and hence the subjective value of a resource. It is shown that linking these ideas with earlier theories on the causation of display components, leads to an explanation of why birds have a greater repertoire of signals associated with aggression/winning, than with submission/losing. It is suggested that modellers of communication systems and those interested in the theory of signal design should pay more attention to the evolutionary constraints imposed by signal origin. Copyright 1999 Academic Press.     

Co-option of male courtship signals from aggressive display in bowerbirds

The pre-existing trait hypothesis suggests that females evolve a mating preference for an already existing male trait. This hypothesis poses a simple resolution to Darwin's long-standing question of how elaborate, male display traits evolve. The frequently observed convergence of aggressive and courtship displays across a wide array of species provides the only current support for this hypothesis. Here we provide much more detailed supporting evidence from bowerbird skrraa calls used in aggression and courtship. Consistent with the pre-existing trait hypothesis we show that (i) putatively co-opted skrraa calls used in courtship and aggression are homologous, (ii) skrraa calls were used in aggression in bowerbirds before being used in courtship, (iii) historically, intense, aggressive-like courtship calls were present near the time of co-option, and (iv) bower types contemporaneous with co-option emphasize design features that provide females protection from the adverse effects of intense courtship displays. These results, plus evidence for a female preference for males with intense aggressive-like courtship skrraa calls, suggest that aggressive skrraa calls have been co-opted for use in male courtship display     

Interference with ritualized threat behaviour in Cichlasoma nigrofasciatum

According to ethological theory, ritualized threat behaviour functions to prevent overt aggression. This prediction was tested in twenty-three pairs of male Cichalasoma nigrofasciatum. Pairs prevented from giving frontal displays by surgically restrained gill covers showed significantly less mouth fighting than sham operates. Pairs prevented from giving frontals and impaired in delivering tail beats because of separated caudal-fin rays showed intermediate aggressive behaviour. Experimental animals in both groups fought and frequently became dominant when paired with sham operates. It was concluded that threat behaviour in C. nigrofasciatum increases subsequent aggressive behaviours both quantitatively and qualitatively. It is suggested that ethological theory has identified the effects of ritualized aggression as a complex behavioural sequence ending in submissive behaviour with that of its first phase alone.     

Estimating the energetic cost of fighting in shore crabs by noninvasive monitoring of heartbeat rate

After establishing shore crabs, Carcinus maenas, individually in separate aquaria, we used a noninvasive infrared phototransducer to monitor their heartbeat rate continously before, during and after fights with intruder crabs. We confirmed that heartbeat rate is a reliable indicator of oxygen consumption and then used it to estimate indirectly the energetic cost of fights differing in duration and intensity, and its dependence on prior residence and relative size of opponent. Prior residence in aquaria significantly increased the probability that crabs would initiate fights against intruders. The majority of fights were resolved by aggressive contacts, display being used extensively only against smaller intruders. Fights between evenly sized opponents and between residents and larger intruders involved almost continuous aggression, whereas fights with smaller intruders involved several shorter bouts of aggression. Fight duration was weakly correlated with the relative size of opponents. Heartbeat rate, measured only in residents, was elevated above resting levels throughout fights, hence energy expenditure during fighting increased linearly with fight duration. Contrary to expectation, heartbeat rate was not significantly influenced by relative size of the opponent or by the intensity of aggression. After fighting, heartbeat rate usually returned to resting levels within 30-60 min, recovery taking longer in fights against larger intruders, when the fight was always lost. We propose that prolonged elevation of heartbeat rate in residents that had lost to larger intruders represented a state of alertness, adaptive against impending risks of resource loss or injury. Copyright 2000 The Association for the Study of Animal Behaviour.     

Factors influencing fighting and threat in the parrot genus Trichoglossus

Parrots of the genus Trichoglossus perform elaborate repertoires of visual threat displays during agonistic interactions. The displays of different species and subspecies differ in complexity and composition. As repertoire complexity increases, taxa tend to emphasize the more exaggerated components of display, and they also show a reduced inclination to attack a mirror-image opponent. These covarying attributes of threat displays in Trichoglossus are highly correlated with interspecific differences in bill-length. These findings are discussed in the light of recent theoretical advances in the study of animal threat and fighting behaviour.     

False Alarm Calls as a Means of Resource Usurpation in the Great Tit Parus major

Great tits Parus major regularly gave alarm calls in winter without the presence of actual or potential predators. Such false alarm calls were given to deceive both conspecifics and heterospecifics. False calls were given when flock-feeding sparrows Passer spp. monopolized a concentrated food source; if the food resource was dispersed false alarm calls were not used, independently of the presence of sparrows. Dominant great tits used alarm calls deceptively if a dominant conspecific was present on a concentrated food source but not if a subdominant individual was present; subdominant great tits were displaced by means of threat displays. Subdominant individuals gave false alarm calls both if dominant or subdominant conspecifics were present. False alarm calls were especially used when food was scarce or when great tits were feeding at a high rate, i.e. during snow storms and in the morning and the afternoon.     

Probing aggressive motivation in a cichlid fish

The duration of startles provides an inverse measure of motivation to resume the previous activity. Here, we use a novel method in which one convict cichlid fish (Amatitlania nigrofasciata) of a competing pair was startled independently of the opponent. Fish were given various opponents and the mean startle duration determined. This mean was negatively correlated with the mean use of highly escalated ‘frontal activities’ such as biting and frontal display, but not the less escalated lateral displays or tail beating. Thus the startle duration was a reliable surrogate measure of the most escalated components of aggressive interactions. That is, it provided a motivational probe for aggressiveness of individual fish. Fight motivation is often determined in terms of fight duration or physiological costs for losers, who reveal the costs they are prepared to pay. We discuss various potential advantages of the motivational probe over previous measures, particularly with respect to winners and losers and different times during the interactions.     

Dominance and aggression in social groups of male and female rats

Two experiments were performed to examine aggression and dominance in domestic male and female $\text{Rattus norvegicus}$ living in small mixed-sex (3 males and 3 females) groups. Experiment 1 examined the development of aggression in females. A single female (alpha) within each of the six colonies tested showed the preponderance of attacks on male intruders placed into the home-cage when male colony residents were absent. Over 12 weeks of intruder-aggression training female alphas showed only a mild nonsignificant elevation of aggressive behavior. A comparison of aggression of male and female colony alphas tested with opponents of each sex revealed that aggression was mainly directed at like-sex opponents, and that female attack was more defensive in character than male attack regardless of opponent sex. The highest intensity of aggression occurred when male alphas confronted male intruders. Although intruders never showed offense toward male residents, 61% of intruding males showed offense in response to attack by females.Experiment 2 investigated the relationship between aggressive dominance and competitive measures of dominance within each of 10 mixed-sex colonies. Alpha stat s of male and female colony residents did not reliably predict priority of access to food or water in tests of direct resource competition with like-sex colony members. When colony males were simultaneously tested for copulation, the copulatory behavior of alpha males was significantly greater than that of other colony males. Results are discussed in relation to the role of aggression in the reproductive strategy of male and female $\text{Rattus norvegicus}$.     

Resolution of aggressive conflicts by immature long-tailed macaques Macaca fascicularis

The behaviour following aggressive interactions differed from the behaviour following friendly or neutral control interactions among unrelated immature male Macaca fascicularis. Former opponents interacted again sooner and more often after aggression than after a friendly or neutral encounter. The higher rate did not reflect a general increase in social activity. The behaviour characterizing encounters between former opponents also differed in the two situations, with more affiliative interactions postconflict. In both situations, however, most interactions between former opponents were initiated by the higher-ranking aggressor. This pattern of interactions between former opponents is interpreted as a repair mechanism for relationships strained by aggression (i.e. reconciliation). A similar pattern of interactions was found for related animals, though it occurred more rarely than among non-kin.