Dominance hierarchies in group-living cleaning gobies: causes and foraging consequences

In species living in social groups, aggression among individuals to gain access to limiting resources can lead to the formation of stable social hierarchies. We tested whether dominance rank in social groups of sponge-dwelling cleaning gobies Elacatinus prochilos in Barbados was determined by physical attributes of individuals or by prior experience of dominance, and examined the foraging consequences of dominance rank. Intraspecific aggression within groups resulted in stable dominance hierarchies that were strongly correlated with fish length. Dominant individuals maintained exclusive territories while subordinate fish occupied broader home ranges. Larger, competitively dominant fish were able to monopolize areas inside the sponge lumen with the highest abundance of the polychaete Haplosyllis spp., a favoured prey item, and achieved the highest foraging rates. The removal of a territorial individual from large groups resulted in a domino-like effect in territory relocation of the remaining fish as individuals moved to the territory previously occupied by the individual just above them in the group hierarchy. Individuals added to existing groups generally failed to gain access to territories, despite being formerly dominant in their original groups. When given the opportunity to choose a location in the absence of larger competitors, gobies frequently preferred positions that were previously defended and that had abundant food. These results suggest that intraspecific competition for resources creates the observed dominance structures and provides support for the role of individual physical attributes in the formation and maintenance of dominance hierarchies.     

The physiology of rainbow trout in social hierarchies: two ways of looking at the same data

Salmonids form dominance hierarchies in environments, where space or food are limiting. Our first objective was to investigate the physiology of individual rainbow trout in 4-fish hierarchies. Our second was to compare conclusions drawn from grouping physiological data on the basis of social rank with those based on relating individual physiology to individual aggressive behavior. To create a social hierarchy, groups of 4 juvenile trout were fed (1 % ration) using a darkened feeding container, twice daily (morning and evening). Each morning feeding was videotaped to record aggressive behavior, thereby facilitating the assignment of a social status rank to each fish. On days 5 and 10–11, physiological parameters were measured in fish fasted for 24 h. Social hierarchies formed in all tested groups. One fish would become dominant, whereas the three subordinate individuals would each assume a stable social rank. When classified according to this social rank, the three subordinate individuals all displayed similar physiology, different from the physiology of the dominant fish. The latter included higher ammonia excretion rate, greater protein utilization in aerobic metabolism, greater feeding, higher specific growth rate, greater increase in condition factor, and lower routine oxygen consumption rate. However, when individual aggression was taken into account, a continuous gradient was observed between aggression and physiology for most parameters, regardless of social status. These relationships could be improved by normalizing the aggression score to the overall level of aggression in each hierarchy. We argue that individual behavior should be considered instead of just social rank when studying the physiology of trout in social hierarchies.     

Diel activity and home range size in relation to food supply in a drift-feeding stream fish

Individuals within a species often compete for resources inboth space and time. In dominance hierarchies individuals withthe greatest competitive ability will occupy prime locationsduring optimal periods. We compared spatial and temporal useof habitat, aggression, and feeding success among giant kokopu(Galaxias argenteus) in dominance hierarchies under normal andreduced food supply. Under normal food supply conditions dominantfish were predominantly nocturnal and maintained large homeranges at night. Conversely, subdominant fish were diurnal andoccupied large home ranges by day but were generally not observedat night. When the food supply was limited, the utilizationof pools was determined by social rank. Dominant fish from eachpool increased diurnal activity, home range size, aggressiveness,and the capture of food items offered by day while simultaneouslyreducing the spatial and temporal activity and habitat use ofsubdominant fish. These results indicate that behavioral changesin large dominant fish influence and reduce the amount of resourcesavailable to subdominant fish.     

Effects of kinship on aggression and RNA content in juvenile Arctic charr

The behaviour of juvenile Arctic charr Salvelinus alpinus was studied in groups of four siblings composed of familiar and unfamiliar individuals or in mixed groups of four where both siblings and unrelated individuals were present. The frequency of aggressive acts was significantly higher in the mixed groups compared to the pure sibling groups and the difference was present at all levels of the hierarchy rank order, based on a dominance index, except the lowest ranked individuals. The difference was significant after but not before feeding, implying that competition with non-kin for a food resource increased the aggression. No significant difference in weight gain was observed between sibling and mixed groups during the 6 days of the experiment, but the RNA contents of lateral musculature in dominant individuals from sibling groups were significantly higher than the corresponding fish in the mixed group, suggesting a difference in growth rate when the experiments ended. No significant difference in RNA content was observed between subordinate fish of the two treatments, i.e. siblings v . mixed.     

State-dependent feed acquisition among two strains of hatchery-reared Arctic charr

The effects of genetic and environmental factors on aggression and feeding hierarchies were studied using X-radiography to measure food intake by hatchery-reared individuals of two strains (Hammerfest and Svalbard) of Arctic charr Salvelinus alpinus . A reduction in food rations and/or water current speed increased intraspecific aggression, and both factors led to increased interindividual variability in food intake, increasing the coefficient of variation (CV). Following a return to pre-manipulation conditions, CVs decreased to their original level. In control groups, CVs and share of group meals were stable throughout the experiment. The increase in CVs following manipulation was the result of a small number of dominant individuals obtaining a high share of the meal. Restriction in food ration affected share of meals, specific growth rates and the frequencies of non-feeding fish, while reductions in water current speed affected only share of meals. Feeding hierarchies were size-dependent in the control groups. In contrast, no relationships between body weight and feeding rank were evident in groups in which food ration or water current speed were reduced. A small, but consistent, difference was revealed in feeding hierarchy responses between the two strains.     

Impact of Environmental Disturbance on the Stability and Benefits of Individual Status within Dominance Hierarchies

Changes in environmental conditions affect social interactions and thus may modify an individual's competitive ability within a social group. We subjected three‐spined sticklebacks, Gasterosteus aculeatus, housed in groups of four individuals, to environmental perturbations to assess the impact on dominance hierarchy stability. Hierarchy stability decreased during increased turbulence or lowered water levels (‘simulated drought’) whereas control hierarchies became more stable in a constant environment. The dominant individual either became more aggressive and remained dominant during the environmental manipulation or was usurped by a lower rank member. Only simulated drought affected rates of aggression where levels of aggression were higher after the water level was dropped which may be the result of an increased encounter rate in these conditions. When there were large size differences between the group members, the dominant individual performed the greatest amount of aggression and ate the largest proportion of food and there was little aggressive behaviour from the lower ranks. In groups of similar‐sized individuals, aggression was much higher. The benefit of being dominant was to gain weight over the experimental period whereas ranks 2 and 3 lost weight. The lowest rank, 4, actually gained weight over the experimental period. This study suggests that it would benefit an individual to be dominant, highly aggressive and gain weight or be submissive, avoid aggressive interactions and, by sneakily obtaining access to food, also gain weight. Altering environmental conditions has a profound effect on social behaviour in this study.     

The influence of social rank in the angelfish,Pterophyllum scalare,on locomotor and feeding activities in a novel environment

To investigate differences in behaviour associated with social rank and environmental conditions, a comparison was made between swimming and feeding activities of dominant and subordinate angelfish, housed in groups of six, with those of angelfish housed singly in identical laboratory aquaria. Subordinate individuals were less active, less ready to feed and consumed less food items than dominants, but their feeding and activity levels were still greater than those displayed by isolated fish used as controls. When fish from each of the above categories were transferred to a novel, identical tank to be tested individually for a 6-day period, little change was observed in previously isolated fish. In contrast, the previous social experience had a marked influence on the behaviour of the other individuals, the effects being related to the social status. The strongest initial response to the new environment was shown by subordinate individuals, with a significant increase in swimming and a significant decrease in feeding compared to the group situation. Subordinates were significantly more active than dominants, who in turn moved more than previously isolated fish. Dominants and subordinates were now similarly reluctant to feed, and their food consumption was less than that of previously isolated fish. With time in the novel environment a significant reduction of swimming activity and a recovery of feeding measures were detected, but levels were still depressed in relation to the group condition, and lower in subordinates than in dominants, indicating the long-lasting effects of the previous social interactions. The results have clear implications for laboratory studies using groups of fish in which social hierarchies may be established prior to individuals being tested singly in a novel environment.     

The effect of ration level and social rank on the development of fin damage in juvenile rainbow trout

The development and severity of fin damage was examined in groups of juvenile rainbow trout Oncorhynchus mykiss of different strength of feeding hierarchies. The development of dorsal and caudal fin damage over time was compared between four groups fed different ration levels (0·25, 0·5, 1·0 and 1·5% body weight day⁻¹) and between individuals of different feeding rank within each group. Dominance hierarchies were assessed from repeated daily measurements of food consumption of individuals using radiography. The feeding and growth data indicated that the strength of the social hierarchy weakened with increasing ration. Caudal fin damage developed with time in all groups whereas dorsal fin damage developed only under limited rations. The severity of both dorsal and caudal fin damage was significantly dependent on the ration size fed to the group, with lower ration groups sustaining more fin damage. The severity of dorsal fin erosion was greater than for the caudal fin. Within the two lower ration groups, subordinate fish suffered the most dorsal fin damage. The results suggested that the severity of dorsal fin damage within groups of juvenile rainbow trout can be used as an indicator of hierarchy Strength.     

The effect of feeding hierarchy on individual variability in daily feeding of rainbow trout, Oncorhynchus mykiss (Walbaum)

Three groups of rainbow trout, Oncorhynchus mykiss , (initial weight 40 g) were fed one of three rations (low, medium or high) for 73 days. Consumption by individual fish within the three ration groups was measured on four occasions (days 27, 55, 64 and 72) using radiography. Food intake by individual fish varied between days and this variability was expressed using the coefficient of variation (CV). Dominant fish within each ration group were defined as individuals with the greater share of the group meal and these fish had low CVs for food intake indicating relatively little variation in daily consumption. By contrast, the fish which had consumed a low mean proportion of the group meal displayed high CVs for food intake, indicating considerable variability in the sizes of individual meals consumed by these fish. As group ration increased, the range of meal sizes and individual CVs in daily feeding decreased, suggesting that the strength of the feeding hierarchy and the variability in individual consumption decreased as food availability increased. It is suggested that radiography can be used to assess social relationships within groups of fish allowing the assessment of feeding hierarchies in larger groups of fish than would be possible by observational techniques.     

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.     

The relationship between dominance rank and feed intake following the introduction of novel feeds to African catfish

The experiment aimed to examine the effect of changing feeds on individual feed intake and feeding and dominance ranks in groups of African catfish. Following feeding on a commercial control feed (Con.) groups ( n  = 3) of 6 catfish were either fed fish meal (FM42) or maize gluten (MG32) based feeds for five days before being switched to the other feed for five days. Feed intake was significantly lower on FM42 than on Con. or MG32 and occurred whether FM42 was fed first or second after Con. Thus, the effect of changing the feed was feed specific. Stability (Kendall's coefficient of concordance) of feeding rank was higher in 5 out of the 6 groups when they were fed MG35 than when fed FM42. Six types of agonistic behaviours were identified and used to assign dominance rank, there were no correlations between dominance and feeding ranks in groups. This was probably due to non‐linear hierarchies with one dominant fish in each group. There were significant correlations between dominance and feeding ranks for combined data over the initial period ( r  = −0·51, P  < 0·005, n  = 36) and when analyzed for the two feeds. Correlations were stronger for MG35 ( r  = −0·41, P  < 0·01) than for FM42 ( r  = −0·33, P  < 0·05). The experiment demonstrated that the introduction of a novel feed can, but does not necessarily, alter feed intake. Of greater significance is the indication that when feed intake decreased the feeding hierarchy and its relationship to dominance rank became less stable.     

The relative influence of prior residency and dominance on the early feeding behaviour of juvenile Atlantic salmon

In many species where social hierarchy mediates conflict over resources, dominant individuals monopolize food, shelter and reproductive opportunities. The benefits of social dominance, however, can often be offset by a prior residence advantage, whereby individuals arriving first in a new habitat obtain, and subsequently defend, the most profitable sites. We investigated the relative influence of these two factors on the acquisition of feeding territories by juvenile Atlantic salmon,Salmo salar L., by placing groups of six individuals of known dominance rank sequentially into an experimental arena with feeding sites of varying quality. Dominants had an advantage over subordinates in the percentage of time spent in a good-quality feeding site and individual feeding rate. There was also a significant time of arrival effect: those individuals that arrived first in a habitat monopolized the resource. The two effects were of approximately equal strength, so that late-arriving dominant fish had similar success as prior resident subordinates. These results indicate that both dominance and prior residence are important in the feeding behaviour of juvenile Atlantic salmon, and that the two factors can have independent and additive effects.Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

The ghost of social environments past: dominance relationships include current interactions and experience carried over from previous groups

Dominance hierarchies pervade animal societies. Within a static social environment, in which group size and composition are unchanged, an individual's hierarchy rank results from intrinsic (e.g. body size) and extrinsic (e.g. previous experiences) factors. Little is known, however, about how dominance relationships are formed and maintained when group size and composition are dynamic. Using a fusion-fission protocol, we fused groups of previously isolated shore crabs (Carcinus maenas) into larger groups, and then restored groups to their original size and composition. Pre-fusion hierarchies formed independently of individuals' sizes, and were maintained within a static group via winner/loser effects. Post-fusion hierarchies differed from pre-fusion ones; losing fights during fusion led to a decline in an individual's rank between pre- and post-fusion conditions, while spending time being aggressive during fusion led to an improvement in rank. In post-fusion tanks, larger individuals achieved better ranks than smaller individuals. In conclusion, dominance hierarchies in crabs represent a complex combination of intrinsic and extrinsic factors, in which experiences from previous groups can carry over to affect current competitive interactions.     

Fight for your breeding right: hierarchy re-establishment predicts aggression in a social queue

Social aggression is one of the most conspicuous features of animal societies, yet little is known about the causes of individual variation in aggression within social hierarchies. Recent theory suggests that when individuals form queues for breeding, variation in social aggression by non-breeding group members is related to their probability of inheriting breeding status. However, levels of aggression could also vary as a temporary response to changes in the hierarchy, with individuals becoming more aggressive as they ascend in rank, in order to re-establish dominance relationships. Using the group-living fish, Neolamprologus pulcher, we show that subordinates became more aggressive after they ascended in rank. Female ascenders exhibited more rapid increases in aggression than males, and the increased aggression was primarily directed towards group members of adjacent rather than non-adjacent rank, suggesting that social aggression was related to conflict over rank. Elevated aggression by ascenders was not sustained over time, there was no relationship between rank and aggression in stable groups, and aggression given by ascenders was not sex-biased. Together, these results suggest that the need to re-establish dominance relationships following rank ascension is an important determinant of variation in aggression in animal societies.     

Group-Living Herbivores Weigh Up Food Availability and Dominance Status when Making Patch-Joining Decisions

Two key factors that influence the foraging behaviour of group-living herbivores are food availability and individual dominance status. Yet, how the combination of these factors influences the patch-joining decisions of individuals foraging within groups has scarcely been explored. To address this, we focused on the patch-joining decisions of group-living domestic goats (Capra hircus). When individuals were tested against the top four ranked goats of the herd, we found that at patches with low food availability they avoided these dominant patch-holders and only joined subordinates (i.e. costs outweighed benefits). However, as the amount of food increased, the avoidance of the top ranked individuals declined. Specifically, goats shifted and joined the patch of an individual one dominance rank higher than the previous dominant patch holder when the initial quantity of food in the new patch was twice that of the lower ranking individual’s patch (i.e. benefits outweighed costs). In contrast, when individuals chose between patches held by dominant goats, other than the top four ranked goats, and subordinate individuals, we found that they equally joined the dominant and subordinate patch-holders. This joining was irrespective of the dominance gap, absolute rank of the dominant patch-holder, sex or food availability (i.e. benefits outweighed costs). Ultimately, our results highlight that herbivores weigh up the costs and benefits of both food availability and patch-holder dominance status when making patch-joining decisions. Furthermore, as the initial quantity of food increases, food availability becomes more important than dominance with regard to influencing patch-joining decisions.     

Social organization of confined male collared lemmings (Dicrostonyx groenlandicus Traill)

This study describes and quantifies the social organization and agonistic behaviour of adult male collared lemmings (Dicrostonyx groenlandicus) in an indoor enclosure. Five groups of four lemmings were observed during separate 8-day tests. Experimental design permitted an increase in group size while density remained constant. When in groups of two, males established stable dominant-subordinate relationships and, when in groups of four, they formed stable non-linear dominance hierarchies. For either group size, frequency of agonistic behaviour declined over time. Habituation, formation of dominance relationships and spatio-temporal partitioning of space are suggested as possible explanations of this decline. Percentage of initial body weight lost during the experiments varied inversely with social rank. Possible causes of this weight loss are discussed. Social relationships in this study are discussed in light of field observations of male lemming home range distribution.     

Social Dominance Hierarchies and Phenotypic Correlates of Dominance in Captive Groups of the Grey‐cheeked Fulvetta Alcippe morrisonia in Taiwan

Dominance hierarchies usually form quickly among avian foraging groups because they are beneficial to most individuals by reducing conflict. Several characteristics that correlate with dominance rank have been identified in birds, but most of these conclusions rely on studies of temperate species. Hence, we studied whether captive group members of a subtropical species, grey‐cheeked fulvetta Alcippe morrisonia, form social dominance hierarchies when competing for food during the non‐breeding season. We also investigated whether sex, age, body condition and fat score were related to an individual's dominance rank which was established by counting aggressive interactions in six captive groups of nine individuals each. In all groups, linear dominance hierarchies were formed whereby yearlings dominated over adult birds, and individuals with a better body condition were also more dominant, while sex and fat score had no discernable effect. Male yearlings had significantly higher body masses and body condition indices than male adults, while female yearlings had significantly higher body masses, body condition indices and fat scores than female adults. However, there were no significant differences between male and female yearlings or adults for any of these variables. We suggest possible reasons for the dominance of yearlings, such as captive conditions or the higher body weight of yearlings.     

A new case of fish-eating in Japanese macaques: implications for social constraints on the diffusion of feeding innovation

This is the first detailed report of social factors affecting fish-eating in Japanese macaques under natural circumstances. We video-recorded a complete event of fish eating, involving a new fish food species for the monkeys on Koshima island. Following the discovery of a large beached sea bass by a peripheral male, we observed a total of 16 individuals feeding on the fish in turns, and interacting around it. The rank order of access to the fish was mainly explained by the spatial position of group members, whereas dominance determined how long the fish was monopolized. Although limited, the tolerated presence of close-bystanders while feeding was affected by kinship and affiliation. Genealogic data suggested that fish-eating behavior was well maintained in terms of maternal lineages. This report should contribute to a better understanding of how social features may constrain the long-term diffusion of feeding innovations in free-ranging primate groups.     

Dominance hierarchies and factors influencing dominance in the guppy Poecilia reticulata (peters)

Dominance behaviour and factors influencing dominance were investigated in small heterosexual and homosexual groups of guppies (Poecilia reticulata) in the laboratory. Straight line nip-right hierarchies were the most frequent type of dominance relationship. In heterosexual groups separate homosexual hierarchies were formed, and females were the more dominant sex. Dominant males performed significantly more reproductive behaviour than lower rank individuals. Agonistic behaviour appears early in young P. reticulata. Factors involved in determining dominance were size, sex and aggressiveness. Individual or role recognition or both may occur in this species. Advantages of dominance are discussed.     

Interindividual variations in feeding and growth in rainbow trout during restricted feeding and in a subsequent period of compensatory growth

Compensatory growth responses of rainbow trout Oncorhynchus mykiss were studied by examining food intake and growth of individual fish held within groups that were switched between regimes that involved full and restricted feeding. Restricted feeding led to marked interindividual variability in food intake, probably as a result of the establishment of feeding hierarchies. This disparity in food acquisition was reflected in highly heterogeneous growth amongst the fish fed low rations. When fish were transferred from restricted to full rations, they became hyperphagic and displayed high rates of growth. Growth compensation was most marked amongst those fish which had shown the poorest growth during the period of feed restriction. These results suggest that the feeding hierarchies established under feed restriction did not persist, but were rapidly broken down when food became increasingly available, enabling the previously suppressed fish to gain access to food and to display rapid growth.