Aggressive behaviour and dominance relationships of the dark chub,Zacco temmincki with special reference to their individual recognition

Aggressive behaviour and dominance relationships ofZacco temmincki were observed by introducing fish into an enclosed pond. Chase (-flee), lateral display, parallel swim and butt were the principal behavioural patterns in aggressive encounters between fish, while chase, resulting in lateral display by the chased fish was the most common behavioural sequence. Initially, mutual behavioural patterns such as parallel swim and mutual lateral display were most evident among the total aggressive acts although chase became dominant three days after introduction into the pond. The dominance matrix constructed from chase-flee interactions during all observation periods contained many reverse attacks (336 out of 2,740 chases). These reverse attacks did not concentrate upon a specific period and were not site-dependent. Examination of chase-flee interactions and the subsequent behavioural pattern revealed that a chased fish reacted to the chaser either by attacking in turn, or performing lateral display etc. roughly in relation to the dominance rank of the chaser. This result implies that fish recognized each other to a great extent during aggressive encounters. It seems likely that such individual recognition was initiated during the early period when mutual behaviour was most frequent, and that some attacks against the dominance order were caused as a result of revolts rather than mistakes.     

Colour and levels of aggression in the midas cichlid

Previous work established that gold morphs of the Midas cichlid (Cichlasoma citrinellum) dominate normal ones of about the same size. Left unresolved, however, was whether the gold morphs dominate because they are inherently more aggressive or because the gold colour inhibits aggression. The issue was clarified here by comparing levels of aggression within groups of golds only, normals only, and golds plus normals. At first the groups of golds were the least aggressive. But 2 days later the levels of aggression in the other groups fell to about that of the golds. We conclude therefore that the gold colour inhibits attack, but that this effect is only discernible in groups when they are establishing inter-individual relationships disappearing when the groups stabilize. We suggest that gold coloration inhibits attacking by stimulating fear responses.     

Early social status and the development of life-history strategies in Atlantic salmon

Atlantic salmon have a variable life cycle. In good growing conditions, underyearling fish may metamorphose into the migratory smolt phase during their second spring, or delay at least a further year. The strategy adopted by particular fish appears to become fixed during their first summer. This paper examines whether either feeding efficiency or dominance in mid-summer correlates with the life-history strategy adopted. Eighty fish were individually marked and their feeding efficiency (= mean handling time for food items) and dominance rank measured under laboratory conditions in mid-July. Growth rates of the fish were then monitored over the next three months, until developmental strategies became apparent. Discriminant and logistic regression analyses revealed that both dominance rank and size attained by July were independent, significant predictors of future developmental pattern (the age at metamorphosis being correctly predicted on the basis of rank and size in 84% of cases) whereas feeding efficiency had no effect. Thus fish that were dominant or larger two months after first feeding or both had a greater probability of migrating after only one year in freshwater than those more subordinate or smaller or both.     

Effect of water renewal on dominance hierarchy of juvenile Nile tilapia

Nile tilapia social position (Oreochromis niloticus) can be mediated by multiple channels, including chemical communication. Absence of chemical cues in the environment prevents hierarchical settlement among pairs, and enhances time spent in confrontations. The aim of this study was to test the effect of continuously renewed water flow on the establishment of hierarchical dominance in Nile tilapia juveniles. In this condition, a high frequency of attacks and disruption on hierarchical stability were expected because chemical cues for hierarchy maintenance could be washed out. After 3 days in isolation, the fish were paired by standard size but not by sex, and submitted to two conditions: continuously renewed water flow (RENEWED, n = 7) and non-renewed water flow (NONRENEWED, n = 8). The paired fish were placed in an aquarium (40 cm × 30 cm × 40 cm) for 3 h; four 10-min sessions were video-recorded: the first, immediately after the fish were paired and the others 1, 2, and 3 h after pairing. Hierarchy was identified by a dominance index (DI = given attacks/received + given attacks) for each fish. The hierarchical stability was achieved by analyzing the difference between dominant DI and subordinate DI (DI-D). Hierarchy was established in both groups after second session because the DI was significantly higher for one fish of the pair. The frequency of attacks of the dominant fish in RENEWED and NONRENEWED conditions was similar in all observation sessions. The attack frequency by subordinate fish was also similar during the first three sessions (2-h pairing). However, the frequency of attacks by subordinate fish in the RENEWED condition was higher than in the NONRENEWED situation at the fourth observation session (means ± S.E.: RENEWED = 2.83 ± 0.94 × 10 min⁻¹ and NONRENEWED = 0.25 ± 0.16 × 10 min⁻¹; Mann–Whitney, p = 0.04). At this point, a significant reduction of the DI-D was observed (means ± S.E.: RENEWED = 0.70 ± 0.11 and NONRENEWED = 1,00 ± 0.002; Mann–Whitney, p = 0.04). The changes in DI-D were related to more frequent attacks by the subordinated fish in renewed water flow. According to our results, the unsteady agonistic interaction under renewed water flow leads to social instability. Thus, continuous water renewing can wash out relevant chemical substances and therefore disturb the dominance recognition by subordinate fish.     

Social rank in winter flocks of Willow Tits Parus montanus

The social hierarchies in winter of ten flocks of Willow Tits Parus montanus were studied when the birds were foraging naturally and when visiting feeders. All the flocks consisted of one adult mated pair together with two juvenile males and two juvenile females (probably pairs). All flocks studied had a stable composition and the hierarchies remained constant throughout the study period. The hierarchies were linear and unilateral. The adults of each sex dominated the respective juveniles and within each age group the male dominated the female. The dominance relationships between the age and sex groups were not consistent. Although the males dominated all the females in six flocks, in one flock the adult female dominated both the juvenile males, but only one of them in three other flocks. The degree of aggression between flock-members was 0.8 encounters per hour, and males initiated 94% of all attacks. Body-weight explained 77% of the variation in dominance rank. It is suggested that the dominance rank of a male is also a function of his seniority, while the rank of a juvenile female is correlated with the rank of her mate.     

Correlation among dominance status, metabolic rate and otolith size in masu salmon

In spatial competition between individuals, neither fish sex nor body mass affected dominance status in masu salmon Oncorhynchus masou . In contrast, resting metabolic rate ( M _R) was significantly correlated with dominance status, indicating that a high metabolic rate can increase the dominance rank of juvenile salmon. Whole animal growth rate was significantly correlated with M _R, but not with initial body weight. This suggests that the body size of masu salmon is not a cause, but rather a consequence, of dominance status which is closely related to M _R. The increment width between otolith daily rings was also significantly correlated with M _R. Thus, the size of the Otolith may indicate the degree of M _R.     

Patterns of agonistic behaviour, shelter occupation and habitat preference in juvenile Lipophrys pholis, Coryphoblennius galerita and Gobius cobitis

The patterns of agonistic behaviour and substratum preferences were investigated in captive groups of juveniles of Lipophrys pholis (Blenniidae), Coryphoblennius galerita (Blenniidae) and Gobius cobitis (Gobiidae). In monospecific groups, size was an excellent predictor of the rank attained by each individual. In heterospecific groups, both C. galerita and L. pholis were dominant over G. cobitis of similar size, and C. galerita dominated L. pholis . Although G. cobitis showed higher rates of activity and of attacks per minute, and a higher ratio of attacks over threats, all these measures were depressed in the presence of either blenniid. In both blennies, in monospecific groups, the dominance rank of each fish was a good predictor of the time spent inside shelters. G. cobitis occupied preferentially the sandy substratum both in mono- and heterospecific groups. Both blennies, when in monospecific groups, occupied preferentially the rocky substratum, with C. galerita showing the highest level of selectivity. In nature, C. galerita also showed a marked preference for rocky substrata, while G. cobitis was especially abundant in mixed bottoms. L. pholis occupied an intermediate position. In the blenniids studied, competition for access to shelter may be one major functional consequence of agonistic behaviour in non-reproductive contexts.     

Biochemical and hormonal correlates of dominance and social behavior in all-male groups of squirrel monkeys (Saimiri sciureus)

This study examined the relationship between dominance rank and several physiological and behavioral measures in stable, captive, all-male squirrel monkey groups. Four groups, each containing three males, were observed for 12 weeks during the breeding season. Relative dominance ranking among males in each group was based on the direction of agonistic behaviors displayed. For each subject, whole blood serotonin (WBS), plasma testosterone (T), and cortisol (C) were sampled 4 to 6 times over the course of the study. Samples were separated by 1- to 2-week intervals. Each group had a stable linear dominance hierarchy. WBS had small intraanimal variance and was positively related to dominance rank. In contrast, T and C were highly variable within subjects and were unrelated to dominance rank. Among alpha males, concentrations of T and C were positively correlated, and WBS and C were negatively correlated. The effect of dominance rank on the relationship between within-animal fluctuations in WBS, T, and C and behavior was also assessed. In dominant, but not subordinate subjects, intraanimal fluctuations in WBS correlated with agonistic behavior initiated, and fluctuations in C and T correlated with huddling. In beta and gamma males, C was related to agonism received, and in gamma males to food stealing received. Dominance status also affected endocrine response to the stress of capture but not the rate of sneezing. Sneezing was positively correlated with T concentrations irrespective of dominance rank. These results extend the association between WBS and dominance rank previously reported in Old World monkeys to a New World monkey species, support previous suggestions that mean T and C titers are not reflective of dominance rank in well-established groups, and indicate that dominance rank affects adrenocortical response to the stress associated with capture and anesthesia.     

Social life histories: jackdaw dominance increases with age,terminally declines and shortens lifespan

Behaviour may contribute to changes in fitness prospects with age, for example through effects of age-dependent social dominance on resource access. Older individuals often have higher dominance rank, which may reflect a longer lifespan of dominants and/or an increase in social dominance with age. In the latter case, increasing dominance could mitigate physiological senescence. We studied the social careers of free-living jackdaws over a 12 year period, and found that: (i) larger males attained higher ranks, (ii) social rank increased with age within individuals, and (iii) high-ranked individuals had shorter lifespan suggesting that maintaining or achieving high rank and associated benefits comes at a cost. Lastly, (iv) social rank declined substantially in the last year an individual was observed in the colony, and through its effect on resource access this may accelerate senescence. We suggest that behaviour affecting the ability to secure resources is integral to the senescence process via resource effects on somatic state, where behaviour may include not only social dominance, but also learning, memory, perception and (sexual) signalling. Studying behavioural effects on senescence via somatic state may be most effective in the wild, where there is competition for resources, which is usually avoided in laboratory conditions.     

Testosterone and Linear Social Dominance Status in Captive Male Dabbling Ducks in Winter

Dominance hierarchies play an important role in avoidance and/or solving conflicts in gregarious species. In dabbling ducks (Anas species), dominance allows for feeding‐site monopolization in winter quarters where resources are generally limited. In addition, male social rank should theoretically favour access to mates. Dominance rank can be associated with morphological traits, and is often correlated with aggressiveness, a behavioural trait generally related to high testosterone levels. In this study, we investigated the existence of a winter group structure based on dominance relationships and tested for a linear hierarchy, in three species of captive male dabbling ducks (mallard Anas platyrhynchos, pintail A. acuta and wigeon A. penelope). We then analysed the relationship between dominance ranks, morphological parameters and testosterone levels measured in early (Oct.) and mid‐winter (Dec./Jan.). We found that the three male groups of the three species exhibited a linear hierarchy. Testosterone levels differed during winter and between species. Morphologic measurements, body mass and body condition were not correlated with individual dominance ranks, whereas dominant males had higher testosterone levels than subordinates. The slopes of the relationships were similar between species and winter period, but the y‐intercepts differed between species and between early and mid‐winter phases. The linear hierarchy found in the three species indicates that dominance relationships strongly structure dabbling duck groups in winter. Lack of correlation between rank and morphological characters, but correlation of rank with testosterone levels suggests that social rank is more dependent on behavioural traits such as aggressive behaviour. The differences between species and winter periods are discussed in relation to migration and wintering phenology.     

The Maintenance of Stable Dominance Hierarchies and the Pattern of Aggression: Support for the Suppression Hypothesis

Three hypotheses have been proposed to explain the development and maintenance of dominance hierarchies. According to the first hypothesis the dominance hierarchy is a result of the animals fighting once at their first encounter and then using the outcome of that fight to determine the rank order. The second hypothesis proposes that a dominance hierarchy reflects the fighting ability of the individuals in the group at each moment and is therefore relatively fluid with individuals continuously fighting for position. A third hypothesis, the suppression hypothesis, states that the dominance hierarchy is based to a large extent on the outcome of the first fight between the individuals but the dominant animal in each pair continuously attacks the subdominant individuals to condition them to lose in future encounters. We studied six well‐established flocks containing six adult hens each (Gallus gallus domesticus). Five of the flocks had linear hierarchies. The aggression was significantly more often directed towards the next low‐ranking individual. There was a good correlation between rank and comb size (height × width), but no significant correlation between rank and weight, or rank and level of fluctuating asymmetry. There was no significant correlation between levels of aggression and similarity of comb size for individuals of neighboring ranks. Our results tentatively support the suppression hypothesis for the maintenance of dominance hierarchies in the domestic hen.     

Genealogical and cross-genealogical dominance relations in a group of free-ranging rhesus monkeys (Macaca mulatta) on Cayo Santiago

Observations of dominance relations in a large group of rhesus monkeys on Cayo Santiago were carried out over a period of 25 months. Dyadic interactions in which an aggressive gesture in one individual was followed by a submissive gesture in the other were recorded as fights and considered reliable indices of dominance. The analysis revealed the following characteristics: (1) Maternal dominance over female offspring; (2) maternal dominance over male offspring up the age six years at which time the son leaves his natal group or remains in the group and rises in rank over his mother; (3) dominance of older brothers over younger male siblings until the age of five years at which time the younger brother rises in rank; (4) rank reversal between sisters when the younger sister reaches the age of three to four years; (5) brother-sister relative rank dependency on age until the male sibling reaches three to four years at which time he rises in rank; (6) linear dominance relations in the crossgenealogical dominance hierarchy; and (7) linear, but unstable, dominance relations in the adult male hierarchy. With few exceptions, the pattern of genealogical, cross-genealogical, and adult male dominance relations in the group under study was consistent with data reported for a small social group (group F) on Cayo Santiago and for Japanese macaques.     

Merging social hierarchies: Effects on dominance rank in male green swordtail fish (Xiphophorus helleri)

When the same set of individuals are placed in different social contexts, some groups members often experience a change in dominance status. We examined the context-dependence of social status using a group fusion protocol in male green swordtail fish (Xiphophorus helleri). Six individuals were matched for size and separated into two groups of three fish. Each triad established a stable hierarchy after which time the two subgroups were merged into one larger assemblage. The maintenance of within- and between-group rank relationships was examined. Relative rank was preserved within each subgroup across social contexts but we found no evidence that familiarity with dominant animals assists individuals of one subgroup in achieving higher rank (coat-tail effects). Dominant individuals from the pre-fusion groups were significantly likely to obtain high status in the merged group and vice versa for subordinate pre-fusion animals. These results demonstrate that social rank in swordtails is relatively impervious to changes in social context, but we address some deviations from this trend. Small differences in standard length were a significant predictor of the most dominant rank in the post-fusion hierarchy, with the largest animals tending to occupy the alpha position. We discuss our results in terms of the potential factors involved in within- and between-group rank maintenance, including individual recognition, winner and loser effects, or asymmetries in dominance-related characteristics.     

The costs of dominance: testosterone, cortisol and intestinal parasites in wild male chimpanzees

Background

Male members of primate species that form multi-male groups typically invest considerable effort into attaining and maintaining high dominance rank. Aggressive behaviors are frequently employed to acquire and maintain dominance status, and testosterone has been considered the quintessential physiological moderator of such behaviors. Testosterone can alter both neurological and musculoskeletal functions that may potentiate pre-existing patterns of aggression. However, elevated testosterone levels impose several costs, including increased metabolic rates and immunosuppression. Cortisol also limits immune and reproductive functions.

Methods

To improve understanding of the relationships between dominance rank, hormones and infection status in nonhuman primates, we collected and analyzed 67 fecal samples from 22 wild adult male chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda. Samples were analyzed for cortisol and testosterone levels as well as intestinal parasite prevalence and richness. 1,700 hours of observation data were used to determine dominance rank of each animal. We hypothesized that dominance rank would be directly associated with fecal testosterone and cortisol levels and intestinal parasite burden.

Results

Fecal testosterone (but not cortisol) levels were directly associated with dominance rank, and both testosterone and cortisol were directly associated with intestinal parasite richness (number of unique species recovered). Dominance rank was directly associated with helminth (but not protozoan) parasite richness, so that high ranking animals had higher testosterone levels and greater helminth burden.

Conclusions

One preliminary interpretation is that the antagonist pleiotropic effects of androgens and glucocorticoids place a cost on attaining and maintaining high dominance rank in this species. Because of the costs associated with elevated steroid levels, dominance status may be an honest signal of survivorship against helminth parasites.     

Changes to feeding and dominance ranks following the introduction of novel feeds to African catfish

The effect of changing feeds on individual feed intake and feeding and dominance ranks in groups of African catfish Clarias gariepinus was investigated. Following feeding on a commercial feed groups (n = 3) of six African catfish were either fed fish meal (FM42) or maize gluten (MG35) based feeds for 5 days before being switched to the other feed for 5 days. Energy intake was significantly lower on FM42 than on MG35, dry matter intake and protein intake were significantly lower on FM42 than on commercial feed and this occurred whether FM42 was fed first or second. There were no significant differences between intake of MG35 and commercial feed. Thus, the action of changing the feed on its own did not affect feed intake since the decrease was shown to be feed‐specific to FM42. Six types of agonistic behaviours were identified and used to assign dominance rank. There were no correlations between dominance and feeding ranks. This was due to non‐linear hierarchies with one dominant fish in each group. Feeding ranks were more stable when feeding MG35 than FM42. Feeding rank stability (Kendall's coefficient of concordance) was significant in five out of six groups fed MG35 (compared with three out of six fed FM42). Feeding rank stability was higher in five out of the six groups when they were fed MG35 than when the same group was fed FM42. The experiment provided evidence that the introduction of a novel feed can, but does not necessarily, alter feed intake and that feed can influence the stability of feeding ranks.     

Agonistic Interactions and Matrifocal Dominance Rank of Wild Bonobos (Pan paniscus) at Wamba

I studied dominance relations in a wild group of bonobos at Wamba, Democratic Republic of Congo. Although agonistic interactions between males occurred frequently, most of them consisted only of display, and physical attacks were infrequent. Dominance rank order seemed to exist among males, but its linearity is unclear. Dominant males rarely disturbed copulatory behavior by subordinate males. However, high-ranking males usually stayed in the central position of the mixed party and, so, would have more chance of access to estrous females. Among females, older individuals tended to be dominant over younger individuals. However, agonistic interactions between females occurred rather infrequently, and most consisted of displacement without any overt aggressive behavior. Dominance between males and females is unclear, but females tended to have priority of access to food. The close social status between males and females may be related to the prolonged estrus of females and their close aggregation during ranging. Existence of a male's mother in the group and her dominance status among females seemed to influence his dominance rank among males. Young adult males whose mothers were alive in the group tended to have high status. In some cases, change in dominance between high-ranking males was preceded by a corresponding change in dominance between their mothers. As the dominance status of females is similar to that of males, mothers may be able to support their sons to achieve high status, stay in the center of the mixed party, and so have greater access to females, which may maximize the number of descendants of the mothers.     

Towards the integration of social dominance and spatial structure

My aim was to show how individual-oriented (or artificial life) models may provide an integrative background for the development of theories about dominance by including effects of spatial structure. Dominance interactions are thought to serve two different, contrasting functions: acquisition of high rank and reduction of aggression. The model I present consists of a homogeneous virtual world inhabited by artificial agents whose actions are restricted to grouping and dominance interactions in which the effects of winning and losing are self-reinforcing. The two functions are implemented as strategies to initiate dominance interactions and the intensity of aggression and dominance perception (direct or memory based) are varied experimentally. Behaviour is studied by recording the same behavioural units as in real animals. Ranks appear to differentiate more clearly at high than at low intensity of aggression and also more in the case of direct than of memory-based rank perception. Strong differentiation of rank produces a cascade of unexpected effects that differ depending on which function is implemented: for instance, a decline in aggression, spatial centrality of dominants and a correlation between rank and aggression. Insight into the origination of these self-organized patterns leads to new hypotheses for the study of the social behaviour of real animals. Copyright 2000 The Association for the Study of Animal Behaviour.     

Sex differences in the impact of social status on hair cortisol concentrations in rhesus monkeys (Macaca mulatta)

Social status impacts stress in primates, but the direction of the effect differs depending on species, social style, and group stability. This complicates our ability to identify broadly applicable principles for understanding how social status impacts health and fitness. One reason for this is the fact that social status is often measured as linear dominance rank, yet social status is more complex than simply high or low rank. Additionally, most research on social status and health ignores the effects of sex and sex-specific relationships, despite known differences in disease risk, coping strategies, and opposite-sex dominance interactions between males and females in many species. We examine the influence of social status, sex, and opposite-sex interactions on hair cortisol concentrations (HCC) in a well-studied species, rhesus macaques, where the literature predicts low ranking individuals would experience more chronic stress. Animals in three captive, seminaturalistic social groups (N = 252; 71 male) were observed for 6 weeks to obtain metrics of social status (rank and dominance certainty [DC]). DC is a measure of one's fit within the hierarchy. Hair samples were collected from each subject and analyzed for HCC. Generalized linear mixed models were used to examine (a) whether rank, DC, or sex predicted HCC; (b) whether same- or opposite-sex dominance relationships differentially impacted HCC; and (c) whether aggressive interactions initiated or received could explain any observed relationships. Results indicated that DC, not rank, predicted HCC in a sex-specific manner. For males, high HCC were predicted by receiving aggression from or having high DC with other males as well as having low DC with females. For females, only high DC with males predicted high HCC. These results likely relate to sex-specific life history pattern differences in inherited versus earned rank that are tied to female philopatry and male immigration.     

Dominance and its Behavioral Measures in a Captive Group of Bonobos (Pan paniscus)

We investigated the existence of a social dominance hierarchy in the captive group of six adult bonobos at the Planckendael Zoo. We quantified the pattern of dyadic exchange of a number of behaviors to examine to what extent each behavior fits a linear rank order model. Following de Waal (1989), we distinguish three types of dominance: agonistic dominance, competitive ability and formal dominance. Fleeing upon aggression is a good measure of agonistic dominance. The agonistic dominance hierarchy in the study group shows significant and strong linearity. The rank order was: 1. female (22 yr), 2. female (15 yr)., 3. male (23 yr.), 4. female (15 yr.), 5. male (9 yr.), 6. male (10 yr.). As in the wild, the females occupy high ranks. There is prominent but nonexclusive female agonistic dominance. Teeth-baring does not fulfil the criteria of a formal submission signal. Peering is a request for tolerance of proximity. Since its direction within dyads is consistent with that of fleeing interactions, it is a useful additional measure to determine agonistic ranks in bonobos. In competitive situations, the females acquire more food than other group members do. The rank obtained from access to food resources differs from the agonistic rank due to female intrasexual social tolerance, expressed in food sharing. We typify the dominance styles in the group as female intrasexual tolerance and male challenging of rank differences. The agonistic rank order correlates significantly with age and has a strong predictive value for other social behaviors.