Intraspecific agonistic interactions in freely swimming mormyrid fish, Marcusenius macrolepidotus (South African form)

South African bulldogs (Marcusenius macrolepidotus, Mormyridae) generate brief (less than 1 ms) electric organ discharges (EODs), separated by much longer and highly variable inter-discharge intervals (IDIs). The diurnal and nocturnal overt behaviour and electrical activity were studied under various conditions: in isolated fish, in pairs of fish, and in a group of four fish that were kept in a "natural" large aquarium. EODs from up to four individuals were recorded simultaneously and identified. While resting during the day, isolated fish showed a broad inter-individual variability of IDI patterns, with distribution histogram modes ranging from 85.7 ms to 325.8 ms. When foraging during the day, IDI modes were shorter and less variable (36.3–48.3 ms). Behaviour patterns displayed during nocturnal agonistic encounters were retreating, parallel swimming, anti-parallel display, attack, and fleeing/chasing. High-discharge-rate (HD) displays were observed at several stages of these encounters, for example, during anti-parallel display (a period of low overt motor activity), or following attacks. IDI durations as short as 11 ms occurred during HD displays, which followed low-rate inter-HD activity almost without transition. IDI distribution histogram modes when fish showed anti-parallel display were 15.4 ms and 24.8 ms, and 30.0 ms during nocturnal non-agonistic interactions. No overt fighting was observed once a dominance relationship was established. In a large aquarium, an approaching dominant male evoked a simultaneous discharge arrest in a group of three subdominant males. Electronic Publication     

Dominance and effects of strange conspecifics on aggressive interactions in the hermit crab Pagurus longicarpus (say)

Dominance orders in the hermit crab Pagurus longicarpus were observed in the laboratory. Groups of four crabs formed loose dominance hierarchies as determined by repeated display and retreat behaviour. Stronger dominance orders were inversely correlated with the frequency of aggressive interactions. Recognition of individuals, as measured by frequency and intensity of aggressive encounters between familiar crabs and introduced strangers, was not important in maintaining dominance orders. Rather, P. longicarpus recognized the aggressive state of conspecifics, as shown by frequencies of aggressive encounters after individuals of different dominance rankings were exchanged between established hierarchies.     

Do Differences in Conspecific Body Size Induce Social Stress in Domestic Rainbow Trout?

Little is known about the cognitive and subjective experiences of fish that are confined with conspecifics of varying body sizes. Plasma cortisol and several behavioural variables were recorded when a ȁ8mediumȁ9 sized fish had its familiar social group (comprised of medium, like-sized individuals) replaced with a group of fish that were either medium sized, smaller or larger than itself. Interestingly, the medium sized test fish showed very few behavioural responses indicative of stress when exposed to a new social cohort. Fish did not use a particular part of their tank, or water column, nor did they show any significant change in locomotory behaviour. There was no difference in aggressive ȁ8chasingȁ9 behaviour in any of the treatments, however, medium sized fish were frequently chased by their tank-mates when exposed to the ȁ8largeȁ9 fish treatment, and were almost never chased by fish smaller than themselves. Similarly, plasma cortisol concentrations did not differ between fish that were exposed to different size treatment groups, although there was a high increase above baseline levels; this suggests that encounters with unfamiliar fish were stressful, regardless of size.     

Is the parallel walk between competing male fallow deer, Dama dama, a lateral display of individual quality?

During competitive encounters protagonists are expected to use signals of individual quality particularly if there is a risk of injury or death. Lateral presentation of body profile, by which information regarding phenotypic characteristics associated with individual quality are displayed, may represent such a strategy. During aggressive interactions, male fallow deer frequently engage in parallel walking which is assumed to represent a mutual display of quality, as mediated by exposure of the maximal profile of the body or antlers. We examined the context and role of the parallel walk during competitive encounters to investigate whether there was evidence that dyads of competing males were assessing differences in phenotypic characteristics. There was no evidence to support the hypotheses that the parallel walk is a lateral display of body size or weaponry or that its use is associated with a reduced level of escalated or risky behaviours during fighting. Total time spent fighting was not shorter when a parallel walk was present than when there was no parallel walk. The parallel walk was highly associated with fighting and it was more likely to be initiated by the subsequent loser. Furthermore, parallel walking frequently followed bouts of fighting and as such may represent a strategy that permits an animal the opportunity to decide whether to continue fighting. Parallel walking was also associated with a failure to resolve contests in favour of one animal indicating that it may be a means of withdrawing from further fighting without incurring a loss in dominance status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Signals and behavioural responses are not coupled in males: aggression affected by replacement of an evolutionarily lost colour signal

Male Sceloporus virgatus lack the blue abdominal patches which are used during aggressive encounters in other Sceloporus lizards. Herein we report that, despite having lost this signal, males have retained a behavioural response to experimentally restored blue abdominal patches. We tested two adaptive hypotheses: selection acted primarily upon signallers or selection acted upon both signallers and receivers. The first predicts that only the signal is lost and that male interactions should be affected by the restoration of blue patches. The latter predicts that both the signal and behavioural response are lost and the display of the restored blue patches should have no effect on male-male interactions. We compared the behaviour of receivers in paired encounters where one male (signaller) had blue-painted abdominal patches to a set of trials where both males had white-painted abdomens, unmanipulated abdomens or a novel-painted pattern. The receivers of the blue-painted signal were more likely to display submissive behaviour. The receivers in either the unmanipulated, white-painted or novel-painted signal trials were more likely to display neutral behaviour. These results support the hypothesis that receivers have retained a behavioural response and selection has acted primarily on the signaller. We believe this is the first documentation of males responding to an evolutionarily lost signal in conspecific males.     

The effects of surgically-implanted dummy radio transmitters on the behaviour of wild Atlantic salmon smolts

The social behaviour of Atlantic salmon smolts was evaluated during their migration period under controlled conditions in an experimental stream tank. Agonistic behaviour, dominance, distance to nearest neighbour, and distance from substrate were examined pre- and post-surgical implantation of dummy radio transmitters (2.4–4.3% body weight). Smolts were able to quickly equilibrate after transmitter insertion. Social ranking changed in nine of the eleven trials with four fish, and in only one of the five trials with pairs. No significant differences were found (p>0.05) in any of the behavioural parameters measured. Overall, the only effect on smolts surgically implanted with radio transmitters was a large shift in dominance. The presence of an antenna also elicited aggressive attacks from other individuals.     

Balancing the competing requirements of air-breathing and display behaviour during male–male interactions in Siamese fighting fish Betta splendens

Air-breathing fish of the Anabantoidei group meet their metabolic requirements for oxygen through both aerial and aquatic gas exchange. Siamese fighting fish Betta splendens are anabantoids that frequently engage in aggressive male–male interactions which cause significant increases in metabolic rate and oxygen requirements. These interactions involve opercular flaring behaviour that is thought to limit aquatic oxygen uptake, and combines with the increase in metabolic rate to cause an increase in air-breathing behaviour. Air-breathing events interrupt display behaviour and increase risk of predation, raising the question of how Siamese fighting fish manage their oxygen requirements during agonistic encounters. Using open-flow respirometry, we measured rate of oxygen consumption in displaying fish to determine if males increase oxygen uptake per breath to minimise visits to the surface, or increase their reliance on aquatic oxygen uptake. We found that the increased oxygen requirements of Siamese fighting fish during display behaviour were met by increased oxygen uptake from the air with no significant changes in aquatic oxygen uptake. The increased aerial oxygen uptake was achieved almost entirely by an increase in air-breathing frequency. We conclude that limitations imposed by the reduced gill surface area of air-breathing fish restrict the ability of Siamese fighting fish to increase aquatic uptake, and limitations of the air-breathing organ of anabantoids largely restrict their capacity to increase oxygen uptake per breath. The resulting need to increase surfacing frequency during metabolically demanding agonistic encounters has presumably contributed to the evolution of the stereotyped surfacing behaviour seen during male–male interactions, during which one of the fish will lead the other to the surface, and each will take a breath of air.     

Agonistic interactions with asymmetric body size in two adult-age groups of the annual killifish Millerichthys robustus (Miller & Hubbs, 1974)

In this study, the author evaluated two adult age groups of the Mexican rivulus Millerichthys robustus with body size asymmetries to determine the strategies used by an annual killifish during agonistic interactions of different ontogenetic stages. To achieve this goal, the author first characterized the ethogram of agonistic interactions of M. robustus composed of seven behavioural units in males and five behavioural units in females. The author then analysed agonistic interaction strategies used by males and females with body size asymmetries in two groups of different adult ages that represent different ontogenetic stages: (a) just after sexual maturity was reached, at 5 weeks of age, and (b) near natural death, at 24 weeks of age. The agonistic behaviour patterns of M. robustus were compatible with the logic of mutual assessment. Large males had an advantage during their interactions in both age groups, winning all of the encounters. Nonetheless, there was more aggression in 5-week-old fish encounters. In addition, small 24-week-old fish were more aggressive than small 5-week-old fish. These changing strategies may be because of the cost–benefits required during a fight at each ontogenetic stage. In the female encounters, size did not predict winners, as both small and large fish won a similar number of encounters, and some contests remained unresolved regardless of age group. There was a tendency for small females of any age to risk more than males in fights to maintain reproductive fitness.     

Agonistic and reproductive behaviors in males of red hybrid tilapia, Oreochromis niloticus (Linnaeus, 1758) x O. mossambicus (Peters, 1852) (Osteichthyes: Cichlidae).

The red hybrid tilapia, Oreochromis niloticus (Linnaeus, 1758) x O. mossambicus (Peters, 1852) is a fertile hybrid used in the semi-intensive level of fish culture in the Northeast of Brazil. It is a territorial cichlid and is highly aggressive towards conspecifics during the breeding season. The purpose of this study was to investigate and describe the aggressive behaviour displayed by the males of this hybrid in non-reproductive and reproductive contexts. Behavioural observations revealed that aggression displayed by the reproductive males of red hybrid tilapia included threatening, undulation, parallel, lateral and frontal attacks, chasing, escape and submission. Possession of a territory influenced male aggressiveness, which was more intense in their own territory than that observed in a neutral situation. The males built nests, irrespective of female presence. All the behavioural patterns were in accordance with those previously described for one parental species, the Nile tilapia, O. niloticus.     

'Better off alone than in bad company': agonistic colour display in mimetic juveniles of two ephippid species

Comparative field observations of agonistic interactions in juvenile leaf-mimicking Platax orbicularis and Chaetodipterus faber (Ephippidae) were conducted in coastal waters of the Pacific and Atlantic Oceans. Similar agonistic behaviour was observed in the two species, in which individuals stopped displaying their mimetic colouration during encounters with conspecifics, to display conspicuous colours, such as transverse stripes along the body. These events were observed occasionally, almost invariably in individuals of smaller body size. Larger-bodied individuals of both species spent less time in agonistic displays. The absolute size of the fish, however, did not appear to affect the outcome of the encounter, suggesting that dominance is a temporary condition, based on the relative size of the opponents during encounters.     

Wolf predation on moose and roe deer: chase distances and outcome of encounters

We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 1997–2003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 15–19% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.     

Development of agonistic behaviour and vocalization in croaking gouramis

The development of agonistic behaviour and vocalization in the croaking gourami Trichopsis vittata was studied from hatching to sexual maturity (4 months of age). Initial interactions started when fry were 11 days old and consisted of approach and flight in a feeding context. More complex threat patterns appeared during dyadic encounters as fish grew older. Lateral display (spreading of median fins in a lateral position) first occurred during the third week, circling shortly afterwards and pectoral fin beating when fish were 7 weeks old. Rapid pectoral fin beating was first accompanied by sound emission at 8 weeks. Initially, croaking sounds were built up mainly of a series of single pulses, each one produced by one pectoral fin. Later, single pulses gave way to double pulses. Furthermore, pulse period and number of pulses increased, while the dominant frequency of croaks decreased significantly with age. After vocalization was established, frontal display, mouth biting and retreat behaviour occurred at the age of 10 weeks. Initially, young exhibited vertical bars which gave way to dots and horizontal bars at 8 weeks when fish started to vocalize. The order of appearance of behavioural patterns during ontogeny corresponds to the order of appearance in fights between adults. This is the first study demonstrating that the ontogenetic development of social signalling comprises characteristic changes in behaviour, vocalization and coloration in a teleost fish.     

Sex-Specific Aggression and Antipredator Behaviour in Young Brown Trout

Sex differences in adult behaviour are often interpreted as consequences of sexual selection and/or different reproductive roles in males and females. Sex-specific juvenile behaviour, however, has received less attention. Adult brown trout males are more aggressive than females during spawning and juvenile aggression may be genetically correlated with adult aggression in fish. We therefore tested the prediction that immature brown trout males are more aggressive and bolder than immature females. Because previous work has suggested that precocious maturation increases dominance in salmonids, we included precocious males in the study to test the prediction that early sexual maturation increase male aggression and boldness. Aggression and dominance relations were estimated in dyadic contests, whereas boldness was measured as a response to simulated predation risk using a model heron. Independent of maturity state, males initiated more than twice as many agonistic interactions as females in intersexual contests. However, males were not significantly more likely to win these contests than females. The response to a first predator attack did not differ between sex categories, but males reacted less to a second predator attack than females. Sexual maturity did not affect the antipredator response in males. Since there is no evidence from field studies that stream-living immature male and female salmonids differ in growth rate, it appears unlikely that the sex differences demonstrated are behavioural consequences of sex-specific investment in growth. It seems more likely that sex-specific behaviour arises as a correlated response to sexually selected gene actions promoting differential behaviour in adult males and females during reproduction. Alternatively, sex differences may develop gradually during juvenile life, because a gradual developmental program should be less costly than a sudden behavioural change at the onset of sexual maturity.     

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.     

Intragroup behavioural changes and the initiation of sex reversal in a coral reef fish in the laboratory

Observations of the coral reef fish Anthias squamipinnis (Peters) suggest that the behavioural control of female-to-male sex reversal is more complex than a simple dependence on the presence or absence of a male or on simple aspects of aggression or aggressive dominance. When the behavioural interactions of all group members of small, bisexual social groups of this fish were examined before and after male removal, two behavioural features were identified as possible factors controlling the initiation of sex change: the profile for behaviours received from, and the percentage of rushes given to other group members. Males and females each showed characteristic profiles both for behaviours given and for behaviours received. On the basis of the behaviour-given profile I identified two types of female. Both types were capable of changing sex following the removal of the male. Two to four weeks were required by sex-reversing fish for the profiles for behaviour given to change to a completely male form. After male removal, the remaining females began to treat the sex-reversing fish as though she the behavioural and coloration changes of sex reversal in a female.     

Dominance status reversal based on pair-bond formation in the convict cichlid (Amatitlania nigrofasciata)

Dominant individuals have access to higher-quality resource; thus, reversing their dominance status would be important for subordinate individuals. Using the convict cichlid fish (Amatitlania nigrofasciata), this study examines whether forming a pair bond can reverse dominance status. Furthermore, I hypothesize that female convict cichlids will incur more dominance reversals from pair-bond formation than males. Dyadic, same-sex contests were conducted to determine dominant and subordinate individuals. Forced pairing of these individuals based on status was followed by polyadic, between-pair contests. The results indicate that individual dominance status does carry over into between-pair competition. Furthermore, dominance reversals do occur in convict cichlids and occur more frequently in females than in males. In addition, dominant males assist their mates during aggressive encounters, and these assists may account for subordinate females winning against dominant females during polyadic contests.     

Relationships of vervet mothers with sons and daughters from one through three years of age

Social relationships between mothers and juvenile offspring were examined in captive, socially-living vervet monkeys (Cercopithecus aethiops sabaeus) to assess the effects of offspring age and sex, and the mother's dominance rank on behavioural interactions. The results indicate that both high-and low-ranking mothers approach and groom their daughters more than they approach and groom their sons. The frequency of both aggressive behaviour toward offspring and support of offspring in agonistic encounters with other group members is influenced by the mother's dominance rank, but not by offsprin sex. Compared to sons, daughters (particularly daughters of high-ranking females) approach and groom their mothers more often, and support their mothers more often in intra-group aggression. The results are discussed in terms of several predictions from parental investment theory and the concept of mutualism.     

The correlation between subordinate fish eye colour and received attacks: a negative social feedback mechanism for the reduction of aggression during the formation of dominance hierarchies

Eye darkening has been linked to social status in fish. The subordinate's eyes darken, while the eyes of the dominant fish become pale. Although this phenomenon has been described in salmonid fishes and in the African cichlid Nile tilapia Oreochromis niloticus, it is unclear whether eye darkening correlates with a reduction in aggressive behaviour. Thus, we evaluated the link between social status and eye darkening. We evaluated whether the eye colours of subordinate fish correlate with the frequency of received attacks in a neotropical fish, the pearl cichlid Geophagus brasiliensis. We paired pearl cichlids and quantified both the aggressive behaviour and the eye darkening of each fish. As has been described for Nile tilapia and Atlantic salmon, a clear-cut hierarchical relationship formed, where dominance and subordination were associated with pale and dark eye colours, respectively. Initially, eye colour darkening was positively correlated with the frequency of received attacks; however, a negative association occurred following eye darkening, in which the intensity of aggressive interactions decreased. Thus, fish that initially received a high number of attacks signalled subordination more rapidly and intensely (rapid and dramatic eye darkening), thereby inducing a negative social feedback mechanism that led to reduced aggression.     

Effects of previous experience on the agonistic behaviour of male crickets, Gryllus bimaculatus

Male solitary animals frequently enter aggressive interactions with conspecific individuals to protect their territory or to gain access to females. After an agonistic encounter, the loser (subordinate individual) changes its behaviour from aggression to avoidance. We investigated agonistic interactions between pairs of male crickets to understand how dominance is established and maintained. Two na?ve males readily entered into agonistic interactions. Fights escalated in a stereotyped manner and were concluded with the establishment of dominance. If individuals were isolated after the first encounter and placed together 15 minutes later, subordinate crickets tended to avoid any further contact with the former dominant opponent. Moreover, subordinate males also avoided unfamiliar dominant and na?ve opponents. They displayed aggressive behaviour only towards unfamiliar subordinate opponents. This suggests that the subordinate male change their behaviour depending on the dominance status of the opponent. Dominant crickets, in contrast, displayed aggressive behaviour towards familiar as well as unfamiliar opponents. If the interval between the first and second encounter was longer than 30 minutes, the former subordinate male showed aggressive behaviour again. However, if the subordinate cricket was paired with the same opponent three consecutive times within 45 minutes, it avoided the former dominant opponent for up to 6 hours following the third encounter. Our results suggest that the maintenance of dominance in male crickets depends largely on the behavioural change of subordinate individuals. Possible mechanisms to maintain dominance are discussed.     

Aggressive behaviour of underyearling rainbow trout in simulated winter concealment habitat

The behaviour of wild underyearling rainbow trout Oncorhynchus mykiss in concealment habitat in a laboratory stream aquarium at 2 and 6°C was recorded daily with an infrared video camera for 90 min over dawn. Aggressive behaviours (threat nips, nips and chases) were frequent during this time as fish entered concealment habitat. Aggressiveness varied widely among fish groups, with a range of 1 to 45 aggressive acts being initiated during a 90-min filming period. Larger fish initiated most of the encounters and removed a higher proportion of fish from concealment than did smaller aggressors. Thirteen per cent of the aggressive acts resulted in the recipient being completely removed from concealment habitat. The highest combined frequency of aggressive acts was 0.64 per visible fish per 10 min period and occurred at relatively low light levels (300 lx). The data suggest that when fish density is high, such as when habitat is limited, aggressive behaviour in winter may cause some underyearling salmonids to be excluded from concealment.