Comparisons of measures of dominance in the laboratory mouse

Dominance in groups of three male mice was assessed in the following ways: the observation of attack and submissive behaviours, the counting of tail wounds, competitive access to water after deprivation, tube dominance, territorial aggression, and access to female mice. Those mice assessed as dominant following behavioural observation had fewer tail wounds, fought more in the territorial aggression test and showed more interest in sexually receptive females. However, correlations of the measures (and subsequent factor analysis) produced no evidence of an underlying dimension which could be labelled dominance.     

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.     

ON THE DEFINITIONS AND FUNCTIONS OF DOMINANCE AND TERRITORIALITY

1. Dominance/subordinance is a relationship between two individuals in which one defers to the other in contest situations. Each such relationship represents an adaptive compromise for each individual in which the benefits and costs of giving in or not giving in are compared. Familiar associates in groups or neighbours on nearby territories may develop relatively stable dominant-subordinate relationships based on individual recognition. Although the aggressive aspects of dominance are usually emphasized, the less conspicuous actions of the subordinate individual are actually more important in maintaining a stable relationship. 2. In evolutionary terms, dominance essentially equals priority of access to resources in short supply. Usually the subordinate, who would probably lose in combat anyway, is better off to bide its time until better able to compete at another time or another place. Both individuals save time, energy, and the risk of injury by recognizing and abiding by an established dominant-subordinate relationship. 3. Dominance can be either absolute or predictably reversible in different locations or at different times. Of the various forms of dominance behaviour, rank hierarchies and territoriality represent the two extremes of absolute and relative dominance, respectively. A dominance hierarchy is the sum total of the adaptive compromises made between individuals in an aggregation or organized group. Many animals seem to be capable of both absolute and relative dominance, and within species-specific limits the balance may shift toward one or the other. High density, or a decrease in available resources, favours a shift from relative to absolute dominance. Some species may exhibit both simultaneously. Social mammals may have intra-group hierarchies and reciprocal territoriality between groups, while the males of lek species may exhibit ‘polarized territoriality’ by defending small individual territories, with the most dominant males holding the central territories where most of the mating takes place. 4. Territoriality is a form of space-related dominance. Most biologists agree that its most important function is to provide the territory holder with an assured supply of critical resources. Territoriality is selected for only when the individual's genetic fitness is increased because its increased access to resources outweighs the time, energy, and injury costs of territorial behaviour. 5. Territoriality was first defined narrowly as an area from which conspecifics are excluded by overt defence or advertisement. The definition has been variously expanded to include all more or less exclusive areas without regard to possible defence, and finally to include all areas in which the owner is dominant. I define territory as a fixed portion of an individual's or group's range in which it has priority of access to one or more critical resources over others who have priority elsewhere or at another time. This priority of access must be achieved through social interaction. 6. My definition excludes dominance over individual space and moving resources, and includes areas of exclusive use maintained by mutual avoidance. It differs from most other definitions in its explicit recognition of time as a territorial parameter and its rejection of exclusivity and overt defence as necessary components of territorial behaviour. There is an indivisible continuum of degrees of trespass onto territories, and functionally it is priority of access to resources that is important rather than exclusive occupancy. 7. There is a similarly indivisible continuum in the intensity of behaviour needed to achieve priority of access to resources. Deciding whether or not an exclusive area is defended leads to the pointless exercise of trying to decide which cues indicating the owner's presence are conspicuous enough to merit being called defence. Concentrating on overt defence emphasizes the aggressive aspects of territorial behaviour rather than the equally or more important submissive aspects such as passive avoidance.     

Social organization and endocrine profiles of <Emphasis Type="Italic">Australoheros facetus</Emphasis>, an exotic freshwater fish in southern Portugal

Australoheros facetus is a neotropical cichlid and an exotic species in the Guadiana and Odelouca basins (Southern Portugal). In this research, we aimed to characterize the main behavioural patterns and circulating hormones, 17β-estradiol (E₂) for females, and testosterone, 11-ketotestosterone (11KT) and cortisol for both sexes, during the formation of social groups, a crucial step in the life-history of A. facetus. A pair-breeding strategy with territorial behaviour and aggressive interactions was found, with a positive correlation between dominance and size for both sexes. There were no significant differences between non-territorial and territorial individuals in the baseline levels of hormones, but 11KT was higher in males when they became territorial, as E₂ for territorial females, while there was no clear pattern for testosterone. In contrast, cortisol was higher in non-territorial males and correlated negatively with social dominance. These results suggest that size is the main driver for social dominance and that formation of stable hierarchies result in higher circulating 11KT and lowers stress in territorial males. Related to A. facetus’ behaviour, aggressiveness and biparental care may be an advantage during the invasion process of A. facetus, facilitating colonization of new habitats.     

Breeding season influxes and the behaviour of adult male samango monkeys (Cercopithecus mitis albogularis)

Troops comprising a high density population of samango monkeys (Cercopithecus mitis) in Natal province, South Africa, experienced an influx of adult males during the breeding season. Observation of one troop revealed that these males competed with one another and with two resident males for access to receptive females. Although both sexes initiated copulation, attempts to do so were more often successful if female-initiated. Males did not interact with non-receptive females and there were no recorded attempts at infanticide. Male-male interactions were agonistic in the presence of receptive females and neutral at other times. No ritualized displays of dominance and subordinance were seen. The significance of these observations for male reproductive strategies is discussed.     

Mating Systems and Male Size in Australian Hylaeine Bees (Hymenoptera: Colletidae)

The mating systems of seven previously unstudied members of the colletid bee genus Hylaeus Fabricius and one of Hyleoides Smith are described. Male mating tactics can be categorized as territorial (perched males defend flowers or other sites that attract receptive females) or non-territorial (patrolling males search for receptive females at flowering plants). The four species in which some territorial males occur are characterized by: 1. grappling fights among males for preferred perches; 2. territorial control by larger males; 3. the possession of prominent spines or other projections on the venter of the abdomen in larger males; and 4. the occurrence of some males that are as large as, or larger than, the largest females of their species (the ‘large-male phenomenon’). In contrast, the four species that lack territorial males are distinctive in that males: 1. do not engage in grappling contests; 2. lack abdominal weaponry; and 3. are smaller than the largest females of their species. In addition, we searched for the large-male phenomenon in museum collections of four species of Hylaeus that exhibit male abdominal spines (presumed to be the weapons used by territorial individuals) and two other species that lack these attributes (presumed non-territorial patrolling species). The results tend to support the sexual-selection-for-fighting-ability hypothesis, which argues that the evolution of unusually large males is a selective consequence of aggressive male—male competition for access to mates. The limitations of the present data set as a comparative test of this hypothesis are discussed.     

Stress, social behaviour, and secondary sexual traits in a male primate

We examined variation in glucocorticoid levels in the mandrill, a brightly coloured primate species, to identify major social influences on stress hormones, and investigate relationships among glucocorticoid levels, testosterone and secondary sexual ornamentation. We collected a total of 317 fecal samples for 16 adult male mandrills over 13 months, including mating and non-mating periods and periods of both dominance rank stability and instability, and compared fecal glucocorticoid levels with dominance rank, rank stability, presence of receptive females, gastro-intestinal parasite infection, fecal testosterone and facial red coloration. Glucocorticoid levels did not vary systematically with dominance rank, but increased when the dominance hierarchy was unstable, and increased in the presence of receptive females. The relationship between dominance rank and glucocorticoid levels changed direction according to the stability of the dominance hierarchy: glucocorticoid levels were higher in subordinate males under stable conditions, but under conditions of instability higher ranking males had higher glucocorticoid levels. The influence of dominance rank also interacted with the presence of receptive females: glucocorticoids were higher in dominant males than in subordinates, but only during mating periods, suggesting that dominant males are more stressed than subordinates during such periods. These findings support previous studies showing that the relationship between glucocorticoids and dominance rank in male baboons is dependent on the social environment. We also found that males with higher glucocorticoids suffered a higher diversity of gastrointestinal parasite infection, in line with evidence that glucocorticoids suppress the immune system in other species. However, we found no support for the stress-mediated immunocompetence handicap hypothesis for the evolution of condition-dependent ornaments: glucocorticoid and testosterone levels were positively related, rather than the negative relationship predicted by the hypothesis, and we found no relationship between red colour and glucocorticoid levels, suggesting that glucocorticoids do not play a role in translating social conditions or physical health into ornament expression in this species.     

Information on Resource Quality Mediates Aggression between Male Madagascar Hissing Cockroaches, Gromphadorhina portentosa (Dictyoptera: Blaberidae)

Male Madagascar hissing cockroaches, Gromphadorhina portentosa Schaum (Dictyoptera: Blaberidae) have a well‐defined dominance hierarchy that has been assumed to explain the outcome of most competitive interactions. We studied whether males of this species would alter their level of aggression towards unfamiliar rivals as a function of changing resource availability and value – two factors that are key to aggression levels in non‐hierarchical species. We quantified male aggression as three variables (aggressive state – behaviours measured by their duration; aggressive act – behaviours measured by their frequency of occurrence; aggressive latency – the latency to first aggressive behaviour, either state or act) and tested for any context‐specific variation within each by manipulating both territorial status (males were either residents or intruders) and access to mates (female present or absent). Both the presence of a female and territorial status affected male aggression towards rivals as measured by duration of aggressive state. Highest levels of aggression were displayed by residents when a female was present. These results show that inter‐male aggression in G. portentosa is tuned to the immediate expected payoff from fighting, and not exclusively aimed at establishing dominance relationships (which can affect future payoffs).     

The effect of differences in body size on the male territorial system of the fly Dryomyza anilis

Drymyza anilis nales defend carcasses, the oviposition sites for females. On carcasses less than 100 g in weight, a single male establishes a teriitory. Ther sex ratio at carcasses is male-biased. Territorial males are larger than other males on average, and move and attack other males more frequently than non-territorial males do. Large carcasses attract more males than smaller ones, but the female, but the advantage of territorial behaviour decreases with increasing density of males. Copulating males are significantly larger than average males, but smaller than territorial males, suggesting than some non-territorial males have access to females. Males seem to be albe to assess a female's egg load, and to adjust the duration of copulation accordingly.     

Assessment of dominance hierarchy through urine scent marking and its chemical constituents in male blackbuck Antelope cervicapra, a critically endangered species

In ungulates the process of chemical communication by urinary scent marking has been directly related to reproductive dominance, territorial defense and proximity to resources. The differences in the frequency of urine marking and chemical composition of urine of males Antelope cervicapra before, during and after the dominance hierarchy period were assessed. The variations in the urine marking and its chemical profiles of dominant males (n = 9), bachelors (n = 5) and sub-adult males (n = 5) were compared to find out how the dominance hierarchy influences the confined blackbuck herd under semi-natural captive conditions. The frequency of urine marking is significantly higher (p < 0.001) in dominant males. Twenty-eight major constituents were identified in the urine of dominant males (before, during and after the dominance hierarchy period), bachelor and sub-adult males. Among these, three specific compounds namely, 3-hexanone (I), 6-methyl-5-hepten-2-one (II) and 4-methyl-3-heptanone (III) were seen only in dominant males urine during the dominance hierarchy period. Based on the behavioural observation and the unique chemical constituents in the urine, it is concluded that the dominant male scent odor suppresses aggression, scent marking, scent production and territorial patrolling activities of subordinate males, through which the dominant male establish their hierarchy and attains success in reproduction.     

饲养条件下川金丝猴群中主雄的移除和替换对社群行为的影响

在上海野生动物园对一群半散养的川金丝猴进行了为期一年的行为学研究。在此期间,猴群中发生了4起家庭主雄被移除和替代事件和一个雌性群因繁殖需要引入一个成年雄猴的事件,该过程在饲养条件下首次被完整记录。观察发现,繁殖群中,在家庭主雄猴的健康状况良好期间,群中各雄性成员间的社会等级关系相当稳定,很少变动。主雄猴有效地控制着群体的秩序,并严格地看护着其家庭中的雌猴免受其他雄猴的侵扰,而且它也很少对其他低序位雄猴主动攻击。全雄群中则再由一个高序位雄猴控制其他低序位雄猴。疾病、衰老、前主雄猴的存在以及被饲养人员从群中移除进行治疗等都可能引起猴群社会发生很大动荡,尤其是全雄群中各雄猴的社会等级序位发生剧烈改变,甚至发生家庭主雄的替代。在本报道中,人为因素在主雄替代过程中起着主要作用。主雄替代一旦成功,新主雄会把原主雄赶入全雄群,攻击追撵其他低序位雄猴,并彻底与全雄群完全脱离。对于每一个新主雄,家庭中的雌猴对其的接纳表现出明显的选择倾向。雌性性选择可能是野生猴群中新家庭群建立一种内在基础机制,同时提示偷配发生的可能性。     

Behavioral and physiological plasticity: Rapid changes during social ascent in an African cichlid fish

In many vertebrates, reproduction is regulated by social interactions in which dominant males control access to females and food. Subordinate males that displace dominant individuals must rapidly adopt behavioral and physiological traits of the higher rank to gain reproductive success. To understand the process of phenotypic plasticity during social ascent, we analyzed the temporal expression pattern of dominance behaviors and circulating androgen levels when socially-suppressed males of an African cichlid fish Astatotilapia burtoni ascended in status. These experiments tested a prediction of the ‘challenge hypothesis’ that, during periods of social instability, male androgen levels are higher than during socially stable times. We found that socially and reproductively suppressed males perform territorial and reproductive behaviors within minutes of an opportunity to ascend in status, and that animals switch from initial expression of territorial behaviors to more reproductive behaviors during territory establishment. Following this rapid response, social stability may be achieved within 1-3 days of social ascent. Consistent with predictions of the ‘challenge hypothesis’, circulating 11-ketotestosterone (11-KT) levels were elevated within 30 min following social opportunity, coincident with increased aggressive behavior. However, territorial behaviors and serum 11-KT levels were then dissociated by 72 h after social ascent, suggesting either rapid social stability and/or increased physiological potential for androgen production. This behavioral and physiological plasticity in male A. burtoni suggests that perception of social opportunity triggers a suite of quick changes to facilitate rapid transition towards reproductive success, and reveals important features of social ascent not previously recognized.     

Boldness,Dominance, and Territoriality in the Color Polymorphic Tree Lizard,Urosaurus ornatus

Social dynamics in territorial species often reflect underlying variation in aggression and other aspects of social dominance among individuals. In ornate tree lizards (Urosaurus ornatus), males differing in dewlap color differ in social dominance: while blue males are the dominant, aggressive morph and always territorial, yellow males tend to exhibit a less‐aggressive satellite behavioral tactic. However, in habitats with fewer available territorial resources, yellow males defend territories and increase in relative abundance. These observations suggest that consideration of social dominance alone may be insufficient to explain U. ornatus' territorial dynamics in the wild. Here, we tested how both dominance and another important behavioral trait, boldness, contribute to the outcome of territorial disputes in tree lizards. We recorded the territorial behavior of blue and yellow male tree lizards (entered in pairs) in an experimental arena. At the end of each trial, we then approached each male and recorded whether it fled (shy) or not (bold) in response to our approach. As expected, dominant blue males exploited the higher quality perch more often than yellow males. However, when approached by a simulated predator, blue males were more likely to flee than yellow males. Thus, while blue males are more dominant, yellow is likely the bolder morph. As a result, this morph may be better equipped to defend territories in riskier environments. We conclude that although dominance asymmetries may predictably drive initial territorial interactions among competing males, variation in other behaviors (like boldness) may perturb the long‐term outcome of these interactions across variable environments.     

Territorial cooperation and social monogamy: factors affecting intersexual behaviours in pair-living snapping shrimp

Among the factors that may contribute to the evolution of social monogamy are selection for extended mate guarding of females and selection for territorial ‘cooperation’. Many socially monogamous taxa are also territorial, with ‘partners’ sharing a single territory, suggesting that one or both partners may benefit by sharing territorial maintenance. Snapping shrimp (genus Alpheus) are socially monogamous and territorial, living in excavated burrows or with host organisms, with females performing all parental care. The territorial cooperation hypothesis predicts that male and female partners share (1) territorial defence, resulting in a reduction in the risk of eviction from the burrow, (2) burrow construction duties, such that individuals in pairs spend less time in burrow construction relative to solitary individuals, and/or (3) foraging duties, by returning food to the burrow, where it is consumed by both partners. UsingA. angulatus as a model species, a territorial defence experiment revealed that females in pairs were significantly less likely than solitary females to be evicted by female intruders, but males in pairs were not significantly less likely than solitary males to be evicted by male intruders. A subsequent experiment revealed that paired males were significantly less likely to be evicted by an intruding male if paired with sexually receptive females than if paired with nonreceptive females. Another experiment revealed that (1) paired females spent significantly more time in burrow construction than paired males, and (2) both males and females consistently returned food items to the burrow, perhaps incidentally provisioning their mates. These data suggest that social monogamy may have been selected for in part because of the advantages of territorial cooperation, as both males and females are likely to benefit by dividing the labour of territorial defence and maintenance. These tests of the territorial cooperation hypothesis are synthesized with data from tests of the extended mate-guarding hypothesis to place snapping shrimp pairing behaviour into a larger construct incorporating both the influence of ecological pressures (territoriality) and mating interactions between the sexes. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Does agonistic dominance imply feeding priority in lemurs? a study inEulemur fulvus mayottensis

We studied agonistic dominance and priority of access to a limited drinking resource in two captive groups of Eulemur fulvus mayottensis.Gender is not systematically related to agonistic dominance or to drinking priority. The dominance hierarchy obtained in the drinking competition test is correlated with the baseline hierarchy in the absence of induced competition, but not with drinking priority. We analyze results within the framework of recent dicussions on female dominance in Malagasy prosimians.     

Heritable variation for territorial success in two Drosophila melanogaster populations

Male fruitflies from stocks from two localities were artificially selected for defence of a food area against intruding males. Both stocks showed a rapid replicable response to selection over a few generations, indicating considerable genetic variation for territorial success in the base populations. Crosses between lines indicate directional dominance for increased territorial success and no maternal or paternal effects. Selected males escalated relatively more frequently against territory residents than control males, and won relatively more escalated encounters. There was no correlated response in body weight. In the presence of territories, selected males had a higher mating success with inseminated females than control males, but did not differ in mating speed. Indirect selection for territorial success was carried out by allowing flies to mate in the presence of different types of resources. After 21–26 generations, males from lines held in cages with discrete resources where territories could be established had greater territorial success than males from lines held in cages with one large resource. This genetic divergence in response to resource type is consistent with the heritable variation in territorial success and mating success of territorial males.     

Timing of mate-locating by males in relation to female activity in the carpenter beeXylocopa varipuncta (Hymenoptera: Apidae)

Males of the carpenter beeXylocopa varipuncta Patton wait for females to visit them as they hover at landmark territories along ridgelines on some spring afternoons. While hovering, males advertize their presence by releasing a pheromone that attracts passing females. If males have limited time to invest in territorial hovering and signaling, then they should engage in these activities more often at times when mate-searching females are most likely to visit landmark territories. The number of females flying near male territories varies greatly over the course of afternoons and from day to day. Measures of female activity and male territorial activity were highly correlated at one study site, both in terms of changes within afternoons and in terms of day-to-day fluctuations. This result supports the hypothesis that males ofX. varipuncta time their mateattracting behavior to maximize contacts with receptive females.     

Social control of sex change in the Red Sea razorfish Xyrichtys pentadactylus (Teleostei,Labridae)

Synopsis The Red Sea razorfish, Xyrichtys pentadactylus, a territorial haremic labrid with dominance hierarchies within the harems. Theory predicts that primary males (fish developing initially as males) should be rare or nonexistent in haremic territorial species because the larger secondary males (males which have undergone sex and/or color change) limit access to females. Histological examination of gonads of 95 specimens showed that all males are derived from females by sex change (i.e. they are secondary males). During five months of field studies 100% of more than 200 observed matings were pair spawnings — the usual mating practice for monandric (having one type of male) species. Sex change in females was induced by male removal in nature. Isolation of four groups of females in aquaria showed that the largest female in the social group changes sex in the absence of a male, demonstrating that sex change is socially-controlled in this species.     

Reef fishes foraging facilitation behavior: increasing the access to a food resource

Fishes associated in schools acquire adaptive advantages by grouping together, e.g., access to a larger variety of food resources, foraging sites, and protection against potential predators. This work presents the first record of a feeding association between the bucktooth parrotfish, Sparisoma radians and the sailor’s grunt Haemulon parra, on Tamandaré reefs, Southwestern Atlantic. Through this association, S. radians gained access to otherwise unavailable food resources to be found inside territorial damselfish domain, thus characterizing an event of foraging facilitation.