Acoustic threat displays and agonistic behaviour in the red-finned loach <Emphasis Type="Italic">Yasuhikotakia modesta</Emphasis>

Agonistic behaviour and the significance of acoustic threat displays were investigated in juvenile red-finned loaches, Yasuhikotakia modesta. This species produced two different vocalizations during agonistic encounters—clicks and butting sounds. Clicks were produced at some distance from the opponent whereas butting sounds were emitted when one fish touched the other with its mouth. This occurred primarily during circling. Both sound types were short broadband signals with the main energies concentrated at about 230 Hz, but clicks were longer in duration and lower in sound level. Agonistic behaviour usually started when one fish approached the other, spread its fins and produced clicks (threat displays), which was followed by parallel displaying, circling and chasing. All fish approached a mirror quickly and displayed aggressively in a parallel position. The number and duration of the threat displays in front of the mirror image were significantly elevated compared with control experiments (rear of the mirror). When playing back click trains in the presence of a mirror image, loaches vocalized significantly less often than during the silent periods, whereas the amount of lateral displaying remained similar. These data indicate that agonistic sounds reduced acoustic displays in red-finned loaches.     

How boat noise affects an ecologically crucial behaviour: the case of territoriality in <Emphasis Type="Italic">Gobius cruentatus</Emphasis> (Gobiidae)

Gobius cruentatus emit sounds during agonistic interactions. In order to evaluate the effect of boat noise exposure on G. cruentatus territorial behaviour, we played a field-recorded diesel engine boat noise during aggressive encounters between an intruder and a resident fish in a laboratory-controlled tank. We tested two factors: role (resident vs. intruder) and condition (noisy vs. silent); the test animals underwent all the treatments in a round-robin design. Agonistic behavior of the residents was modified by boat noise: during the playback residents were more submissive and won less encounters than in the control (silent) condition. We suggest that sound production is an effective tool for territorial defense, since the impairment of acoustic communication due to the recreational boat noise diminished the ability of the resident to maintain its territory.     

Aggression among sea urchins on Caribbean coral reefs

The existence and nature of intra- and interspecific aggression were examined for five species of sea urchins inhabiting Caribbean coral reefs. The studies took place in the San Blas Islands of Panama and involved the following species: Echinometra lucunter (Linnaeus), E. viridis Agassiz, Diadema antillarum Phillipi, Lytechinus williamsi Chesher, and Eucidaris tribuloides (Lamarck). An intruder was placed next to an undisturbed resident and the behaviors and responses of both individuals were followed. All pairwise combinations of resident species and intruder species were tested except for combinations involving D. antillarum as intruder.

For E. viridis and E. lucunter, agonistic interactions occurred commonly (46–79% of trials) between conspecifics and congeners. Almost all of the agonistic encounters involved pushing. Additionally, biting occurred in 8–25 % of the trials. Residents were most often the aggressors and usually succeeded in retaining their location. Intruders only succeeded in forcing residents out of their positions if the intruder was equal to or larger in size than the resident. Surveys of an undisturbed population of E. viridis during the daytime indicated that 16% of the individuals were engaged in intraspecific agonistic interactions at any one time.

D. antillarum exhibited biting behavior against both species of Echinometra in 23–24% of the trials. Biting attacks against L. williamsi and E. tribuloides occurred rarely. L. williamsi only once demonstrated pushing behavior and never was observed biting another sea urchin. E. tribuloides occasionally exhibited pushing and biting behaviors, both as residents and as intruders, and was twice observed biting Echinometra.

These studies suggest that two kinds of agonistic interactions may commonly occur among Caribbean reef-dwelling sea urchins: (1) intraspecific and interspecific aggression among Echinometra, and (2) predatory/aggressive attacks against Echinometra by D. antillarum. The former may result in greater dispersion of Echinometra relative to food and shelter resources and control the spatial distribution and concentration of Echinometra grazing pressure within an area. The attacks by D. antillarum may result in the restriction to, or higher densities of, Echinometra in crevices and rugose microhabitats that provide shelter from the larger-bodied D. antillarum.     

Analysis of waving and sound‐production display in the ghost crab,Ocypode stimpsoni

The ghost crab Ocypode stimpsoni displays waving and sound production. Sounds are produced by thumping the sand substratum with the major cheliped, and two types of sounds can be discriminated; one with a low frequency of about 12 Hz, called rapping, and another with a higher frequency (about double), called quivering. In our observations, a sequence of waving and sound emission would sometimes terminate abruptly, or appear as independent components but the component order never changed. The most frequently observed patterns were “waving with rapping and quivering”;, “waving with quivering”; and “quivering only”; quivering sounds being involved in more than 80% displays. Quivering sometimes occurred immediately after crabs emerged from the burrow, or when they returned to the entrance after discarding an excavated sand mass. The occurrence frequency of waving and sounds, the wave amplitude, and the frequency of the sound increased when other crabs approached.     

Feature-detecting auditory neurons in the brain of a sound-producing fish

The mormyrid fish Pollimyrus adspersus has auditory specializations for sound pressure detection and uses acoustic displays in its natural social behavior. In this paper it is shown that auditory neurons in the mesencephalon (torus semicircularis) are activated selectively by temporal features of complex sounds. Single neurons were recorded while presenting sounds to fish underwater. The stimuli were acoustic click trains, 400 ms in duration, and were synthesized with differing inter-click-intervals (ICIs). The natural sounds of this species are composed similarly and the range of ICIs synthesized overlapped with the natural range (5–40 ms). One-third of the neurons studied were strongly selective for a narrow range of ICIs, increasing spike rate by ten fold or more at the best ICI compared to the minimum response observed. The best ICI for interval selective neurons remained stable when the sound pressure of the stimulus was changed. Neurons that were selective gave phasic responses to tone bursts, and most had non-monotonic rate level functions. The origin of interval selectivity is discussed and a time-based computational mechanism is proposed. Accepted: 20 December 1996     

Sound production in the Ponto‐Caspian goby Neogobius fluviatilis and acoustic affinities within the Gobius lineage: implications for phylogeny

The aim of this study was to describe the vocal repertoire of the Ponto‐Caspian goby Neogobius fluviatilis and to compare the acoustic properties of this species with those of other soniferous Mediterranean gobies belonging to the Gobius lineage. Vocalizations and associated behaviours were recorded under controlled aquarium conditions in female and male N. fluviatilis. Sound emission was elicited by means of ‘intruder tests’, using an individual of the same or opposite sex as an intruder, and recording sounds using a hydrophone placed 20 cm from the shelter used as a nest for the resident fish. Five acoustic properties, including spectral and temporal properties, were measured from 13 individuals. The vocal repertoire of the species consisted of sequences of short vocalizations during both agonistic and reproductive intraspecific interactions. The wave form of each sound resolved in a pure sine wave composed of rapidly repeated pulses. Sounds lasted about 200 ms, showing an average fundamental frequency of about 80 Hz. Sound properties did not differ between reproductive and the aggressive contexts, and the general structure of sounds was highly stereotyped. The individual means of three acoustic independent traits characterizing the sounds of seven species of the Gobius lineage, including N. fluviatilis, were then entered in a discriminant function analysis to assess how well species could be differentiated on the basis of acoustics, and their degree of affinities. The results suggested that the pulse repetition rate of the sounds, i.e. the relative tonal/pulsatile nature of the sounds, was the most important property in differentiating the species, and that this trait may contain a high level of phylogenetic signal, as the species producing tonal sounds clustered together, in line with the results of recent molecular phylogenic studies. The results were discussed in light of the geological and phylogeographical events believed to have driven the diversification of European gobies.     

Neuroendocrine and behavioral response to social confrontation: residents versus intruders, active versus passive coping styles

We investigated in the present study the neuroendocrine correlates in intruder and resident rats of a social confrontation. Adult male Wistar rats (intruders) were introduced into the home cage of a well-trained resident to induce characteristic agonistic interactions including physical attacks prior to separation by a wire mesh. The hypothalamic-pituitary-adrenal (HPA) axis activity and the intrahypothalamic release of arginine vasopressin (AVP) were monitored via chronically implanted jugular venous catheters and microdialysis probes aimed at the hypothalamic paraventricular nucleus (PVN), respectively. Based on the behavioral data collected during the 30-min confrontation, intruders and residents were additionally classified into two different subgroups: intruders which showed almost no freezing behavior (active copers) versus those showing pronounced freezing behavior (passive copers) and residents which were either predominantly aggressive or non-aggressive. The neuroendocrine data show that social confrontation caused a significantly increased secretion of the adrenocorticotropic hormone (ACTH) into plasma in both intruder subgroups, independently of their coping strategy. In contrast, plasma ACTH in residents was increased in response to social confrontation in non-aggressive animals only, whereas aggressive residents failed to mount an ACTH response. Interestingly, plasma AVP decreased in response to social confrontation in active intruders. As measured in microdialysates, the two groups of residents and passive intruders failed to show significant changes of intra-PVN release of AVP. In contrast, an increased release of this neuropeptide within the PVN could be monitored for active intruders. The data of the present study suggest that the different interpretation of an aversive encounter results in differences in the neuroendocrine response and intrahypothalamic vasopressinergic signaling in intruders versus residents.     

Residency Duration and Shelter Quality Influence Vibratory Signalling Displays in A Territorial Caterpillar

Residents are more likely to win territorial disputes than intruders. One explanation for this prior resident advantage is that residents place a higher value on the resource and are therefore more motivated to win. Although value asymmetry models of animal contests often assume that contestants use information about resource value, information on the proximate cues affecting territorial behaviour is often lacking. We use a simple model system – territorial behaviour in the masked birch caterpillar (Drepana arcuata) to identify factors that affect territorial behaviour. Late instar caterpillars occupy solitary silken leaf shelters, which they defend against wandering conspecifics with a vibratory display. We evaluated how a caterpillar identifies itself as the owner and the factors that influence a resident's motivation to signal. To do so, we conducted three experiments between size‐matched residents and intruders to assess how residency duration and shelter quality independently affected territorial displays during the early stages of a contest. Experiment 1 (Time Exp.) demonstrated that resident signalling rates increase with increased duration on the leaf prior to introducing the intruder. Residents also signal more than intruders after residency periods of 1–3 min and longer, demonstrating that residents gather information about resource value shortly after occupying a leaf. Experiment 2 (Squatter Exp.) aimed to disentangle the effects of time on the leaf and silk accumulation. Squatters (individuals in a shelter made by another) placed for 1–3 min on a leaf containing a full silk shelter signalled more to intruders than did caterpillars placed on a fresh leaf for 1–3 min. Experiment 3 (Shelter Removal Exp.) showed that residents whose shelters had been removed signal less than those occupying an intact shelter, despite an equal length of time investing in them. Our experiment is the first to covary both prior residency duration and territory quality, and we find that the motivation of caterpillars to signal is a function of both of these attributes.     

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.     

Singing above the chorus: cooperative Princess cichlid fish (<Emphasis Type="Italic">Neolamprologus pulcher</Emphasis>) has high pitch

Teleost fishes not only communicate with well-known visual cues, but also olfactory and acoustic signals. Communicating with sound has advantages, as acoustic signals propagate fast, omnidirectionally, around obstacles and over long distances. Heterogeneous environments might favour multimodal communication, especially in socially complex species, as the combination of modalities’ strengths helps overcome their individual limitations. Fishes of the ecologically and morphologically diverse family Cichlidae are known to be vocal. Here we investigated sound production in the socially complex Princess cichlid Neolamprologus pulcher from Lake Tanganyika in East Africa. We show that wild and captive N. pulcher produce only short-duration, broadband high-frequency sounds (mean: 12 kHz), when stimulated by mirror images. The evolutionary reasons for this “low frequency silencing” are still unclear. In laboratory experiments, N. pulcher produced distinct two-pulsed calls mostly, but not exclusively, associated with agonistic displays. Princess cichlids produce these high-frequency sounds both in combination with and independent from visual displays, suggesting that sounds are not a by-product of behavioural displays. Further studies on the hearing abilities of N. pulcher are needed to clarify whether the high-frequency sounds are used in intra- or inter-specific communication.     

DELIMITING SPAWNING AREAS OF WEAKFISH CYNOSCION REGALIS (FAMILY SCIAENIDAE) IN PAMLICO SOUND,NORTH CAROLINA USING PASSIVE HYDROACOUSTIC SURVEYS

ABSTRACT

Exact locations of spawning areas used by marine fishes are needed to design marine reserves and estimate spawning stocks. The location of spawning areas of soniferous fishes such as weakfish Cynoscion regalis can be determined by means of passive hydroacoustic surveys. We conducted nocturnal hydrophone surveys at 12 locations in Pamlico Sound in May of 1996 and 1997. Digital audio tapes were made of weakfish “purring” sounds, the tapes were analyzed spectrographically and compared with ichthyoplankton surveys taken at the same stations and times. All weakfish “purring” sounds were recorded at stations near inlets. Maximum sound pressure levels recorded after sunset were 127 dB (re 1 (μPa) for individual weakfish, but reached a maximum of 147 dB when weakfish and other fish were producing sounds simultaneously. The maximum distance that an individual weakfish “purr” can be detected above the background sound, assuming a cylindrical spreading model, is approximately 50 m. There was a strong association (r = 0.78) between the log₁₀— transformed abundance of early-stage sciaenid-type eggs and maximum sound pressure levels, with the greatest numbers occurring at the inlet stations. These results suggest that passive hydroacoustic surveys can be used to delimit spawning areas for conservation and management purposes.     

Acoustic and visual adaptations to predation risk: a predator affects communication in vocal female fish

Predation is an important ecological constraint that influences communication in animals. Fish respond to predators by adjusting their visual signaling behavior, but the responses in calling behavior in the presence of a visually detected predator are largely unknown. We hypothesize that fish will reduce visual and acoustic signaling including sound levels and avoid escalating fights in the presence of a predator. To test this we investigated dyadic contests in female croaking gouramis (Trichopsis vittata, Osphronemidae) in the presence and absence of a predator (Astronotus ocellatus, Cichlidae) in an adjoining tank. Agonistic behavior in T. vittata consists of lateral (visual) displays, antiparallel circling, and production of croaking sounds and may escalate to frontal displays. We analyzed the number and duration of lateral display bouts, the number, duration, sound pressure level, and dominant frequency of croaking sounds as well as contest outcomes. The number and duration of lateral displays decreased significantly in predator when compared with no-predator trials. Total number of sounds per contest dropped in parallel but no significant changes were observed in sound characteristics. In the presence of a predator, dyadic contests were decided or terminated during lateral displays and never escalated to frontal displays. The gouramis showed approaching behavior toward the predator between lateral displays. This is the first study supporting the hypothesis that predators reduce visual and acoustic signaling in a vocal fish. Sound properties, in contrast, did not change. Decreased signaling and the lack of escalating contests reduce the fish’s conspicuousness and thus predation threat.     

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.     

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”.     

Behavioural responses of male ruffed grouse (Bonasa umbellus,L.) to playbacks of drumming displays

Birds use a variety of sounds in their courtship displays, but the majority of behavioural studies have focused on vocalizations. In contrast, little is known about how non‐vocal sounds, or sonations, are used, even though many avian species produce them. The ruffed grouse (Bonasa umbellus) is a useful species to examine non‐vocal sounds because they lack vocal components in their courtship and rely on a non‐vocal sound to attract mates and defend their territory. Their courtship display, known as “drumming,” is created by the wings, and the number of pulses and speed (pulse rate) varies significantly among males. Anecdotal evidence suggested that males can affect the drumming behaviour of neighbouring males in drumming “duels” in an analogous way to song contests. Here, we test whether males do respond to the playback of drumming sounds of an unfamiliar male. Using a portable speaker system, we played recordings of drumming displays to males that were actively drumming themselves. Throughout each playback, we recorded the drumming behaviour of target males so that we could assess whether drumming activity changes following a playback as well as whether males change the speed of their display. Overall, male grouse were equally likely to approach the speaker or continue drumming following a playback. For those males that continued drumming, their drumming pulse rate was significantly faster following playbacks, but they drummed less often. These results indicate that male ruffed grouse do respond to drumming sounds, but the specific response differs among males. Because the differential response was not related to colour phase or whether a male was drumming in proximity to other males, we suggest that the response of individuals likely varies with other traits, such as hormone levels or behavioural syndrome.     

Behavioral context of echolocation and prey-handling sounds produced by killer whales (Orcinus orca) during pursuit and capture of Pacific salmon (Oncorhynchus spp.)

Availability of preferred salmonid prey and a sufficiently quiet acoustic environment in which to forage are critical to the survival of resident killer whales (Orcinus orca) in the northeastern Pacific. Although piscivorous killer whales rely on echolocation to locate and track prey, the relationship between echolocation, movement, and prey capture during foraging by wild individuals is poorly understood. We used acoustic biologging tags to relate echolocation behavior to prey pursuit and capture during successful feeding dives by fish-eating killer whales in coastal British Columbia, Canada. The significantly higher incidence and rate of echolocation prior to fish captures compared to afterward confirms its importance in prey detection and tracking. Extremely rapid click sequences (buzzes) were produced before or concurrent with captures of salmon at depths typically exceeding 50 m, and were likely used by killer whales for close-range prey targeting, as in other odontocetes. Distinctive crunching and tearing sounds indicative of prey-handling behavior occurred at relatively shallow depths following fish captures, matching concurrent observations that whales surfaced with fish prior to consumption and often shared prey. Buzzes and prey-handling sounds are potentially useful acoustic signals for estimating foraging efficiency and determining if resident killer whales are meeting their energetic requirements.     

Does increased habitat complexity reduce predation and competition in coral reef fish assemblages?

Greater habitat complexity is often associated with a greater abundance and diversity of organisms. High complexity habitats may reduce predation and competition, thereby allowing more individuals to occupy a given area. Using 16 spatially isolated reefs in the Bahamas, I tested whether increased habitat complexity reduced the negative effects of resident predators and competitors on recruitment and survival of a common damselfish. Two levels of habitat complexity were cross-factored with the presence or absence of two guilds of resident fishes: predators (sea basses and moray eels) and interference competitors (large territorial damselfishes). I monitored subsequent recruitment and recruit mortality for 60 days. Residents had strong negative effects on recruitment regardless of habitat complexity. In the presence of residents, recruits suffered high mortality immediately after settlement that was similar on low and high complexity reefs, although high complexity reduced mortality of recruits that survived this early postsettlement period. Comparisons between shelter hole diameters and the sizes of residents suggest that territorial damselfishes and small resident predators could access most shelter holes, whereas large resident predators were excluded from many shelter holes. This study demonstrates that whether habitat complexity reduces predation and competition may depend on several key factors, such as the availability of appropriate shelter, behavioral attributes of interactors, and developmental stage of prey/inferior competitors.     

Vocalization during Agonistic Behaviour in Cottus gobio L. (Cottidae): An Acoustic Threat Display

Agonistic behaviour in the river bullhead C. gobio consists of visual (raising gill covers and fins, lowering the head, darkening) and acoustic (single knock sounds and trains of knock sounds) threat displays, rarely followed by attacks and bites. This study investigates the relationship of vocalizations with size, dominance, territory dimensions and sex of the opponents. Four groups, each consisting of a big male, a small male and a female, were each investigated for three different days. The number of won contests of each individual, the numbers of each sound produced during these encounters and the tank part where encounters took place were determined. Subordinate fish emit fewer sounds but relatively far more trains of knock sounds than dominant ones. They produce relatively more sounds under shelters whereas dominants do this on uncovered areas. α-fish produce more calls during agonistic encounters with β-fish than Ω fish. In β-individuals no such difference was noted. Basically no sex related behaviour could be observed. In each area of the tank one individual won most contests (= territory). Dimensions of territories differed significantly between individuals in each tank (α-, β-, Ω-fish). In no case were all three individuals able to maintain territories. Relative sizes of fish correlate significantly with relative numbers of successful encounters and with territory dimensions. Furthermore, both parameters are positively correlated with the numbers of sounds emitted by an individual. Sound production in C. gobio functions as an acoustical threat display. Because of the high energy costs of sound emission underwater it might be a very effective method of assessing the fighting ability of an opponent.     

Effect of environmental enrichment upon resource holding power in fish in prior residence situations

Resource holding power (RHP), as expressed by gaining dominance, can be affected by extrinsic and intrinsic factors. Extrinsic factors that increase the RHP include e.g. prior exposure to the contest area. The pay-off asymmetry hypothesis was tested according to which there is an asymmetry in the value the resident's territory has for the resident and intruders, i.e. the resident loses more than intruder opponents when losing the contest over a territory in which it has invested much energy during the exploration of food and shelter resources, during the settlement of conflicts with territorial neighbours, etc. This asymmetry will increase the resident's chance to win the fight over intruders. If the resident-intruder asymmetry is enhanced by presenting resource-rich territories to e.g. fish (structurally diverse or 'rich' aquariums), the hypothesis predicts that the resident's dominance chance (probability to win the fight) will be higher than in structurally 'poor' aquariums. It was found that residents were proportionately more often dominant (fight winners) than intruders in the rich compared to the poor aquariums in all seven tested fish species. This demonstrates that a high territory resource value (aquarium enrichment) significantly facilitates the expression of the resident's dominance advantage (prior residence effect). In contrast to dominance, aggressive behaviour before the dominance settlement did not generally differ between residents and intruders when compared in rich and poor aquariums. This suggests that dominance in itself and aggression prior to dominance settlement are at least partially guided by different motivational systems.     

Inter-male aggression with regard to polygynous mating system in Pampean grassland mouse, Akodon azarae (Cricetidae: Sigmodontinae)

Based on the hypothesis that, in Akodon azarae, polygyny operates through female defence, we studied inter-male aggression in order to test the following predictions: during the breeding period (1) resident males are more aggressive than intruder males in the presence of females (FP), and (2) aggressive behaviour is independent of male condition (resident or intruder) in the absence of females (FA). To test our predictions, we used the resident male behavioural response towards an intruder male in relation to FP or FA. We conducted 30 encounters in FP and 27 in FA in 0.79-m² round enclosures placed in the Espinal Reservation. Our results support the prediction that, in FP, the intensity of aggressive behaviour exhibited by males varied in relation to resident or intruder condition. Resident males showed high levels of aggression towards intruders, and intruders exhibited the greatest values of submissive behaviours with residents. In FA, the intensity of aggressive behaviour did not vary in relation to resident or intruder condition. Both resident and intruder males exhibited low aggressive behaviour and inter-male encounters resulted mainly in non interactive behaviours. Our results support the hypothesis that, in A. azarae, the polygynous mating system operates through female defence.