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.     

Dominance and huddling behaviour in Bronze Mannikin Lonchura cucullata flocks

Bronze Mannikins Lonchura cucullata form flocks of up to 30 birds at the end of the breeding season and huddle in communal nests during winter. Assuming there are thermoregulatory benefits associated with huddling for these small endotherms, particularly during winter, we predicted that dominant individuals in flocks would obtain the central locations within the huddles. We tested this prediction by determining the dominance hierarchies of six captive flocks of five individuals by examining interactions at a feeder. Mannikins were observed to huddle when cold-stressed. Although mannikins do not naturally form linear huddles, for ease of observation we recorded the positions of individually marked birds in linear huddles at 5 °C before and after disturbances every half hour for 3 h. Captive flocks of mannikins established linear dominance hierarchies and dominant individuals obtained the central location within the huddle. The central position in the huddle may be beneficial in thermodynamically challenging environments through reduced energy expenditure.     

Females competing to reproduce: dominance matters but testosterone may not

The associations among aggression, testosterone (T), and reproductive success have been well studied, particularly in male birds. In many species, males challenged with simulated or real territorial intrusions increase T and levels of aggression, outcomes linked to higher dominance status and greater reproductive success. For females, the patterns are less clear. Females behave aggressively towards one another, and in some species, females respond to a social challenge with increases in T, but in other species they do not. Prior work on female dark-eyed juncos (Junco hyemalis) had shown that experimental elevation of T increases social status and intrasexual aggression. Here, we conducted two experiments designed to answer three questions: Are endogenous concentrations of T associated with dominance status in captive female juncos? Does dominance status influence readiness to breed in female juncos? And do captive females increase T in response to a challenge? In the first experiment, we introduced two females to a breeding aviary, allowed them to form a dominance relationship and then introduced a male. We found that dominant females were more likely to breed than subordinates, but that dominance status was not predicted by circulating T. In the second experiment, we allowed a resident male and female to establish ownership of a breeding aviary (territory) then introduced a second, intruder female. We found that resident females were aggressive towards and dominant over intruders, but T did not increase during aggressive interactions. We suggest that during the breeding season, intrasexual aggression between females may influence reproductive success, but not be dependent upon fluctuations in T. Selection may have favored independence of aggression from T because high concentrations of T could interfere with normal ovulation or produce detrimental maternal effects.     

Urine-marking during agonistic encounters in the bush dog (Speothos venaticus)

During staged encounters, bush dogs of the same sex showed a high level of aggressive-defensive behaviors and a higher than normal frequency of urine-marking. Where dominance was established during encounters, dominant individuals marked more than subordinates. Where dominance was not clearly established, both participants remained aggressive and showed higher than normal levels of marking. Urine-marking appears to communicate aggressiveness and may be used to compensate for the otherwise relatively inconspicuous aggressive and dominance displays of this species.     

Social Control and Physiological Cost of Cheating in Status Signalling Male House Sparrows (Passer domesticus)

Flock-forming passerines often use plumage characteristics to signal their social dominance. While the benefits to signal dominance seem obvious, costs associated with status signalling are ambiguous. The social control hypothesis predicts that individuals of high social status – with large badges – are involved in more social interactions with individuals of similar badge size. Cheaters are therefore exposed to increased risk of fighting with high quality individuals and the costs associated with enhanced fights with dominant males are supposed to outweigh the benefits of cheating. We tested the social control hypothesis in male house sparrows ( Passer domesticus ), by observing social interactions in captive flocks and determining dominance relationships. Two low status individuals within each flock had the size of their badge experimentally increased and the interactions involving experimental and control birds were recorded. We also assessed the potential physiological cost of cheating in terms of enhanced levels of the stress hormone, corticosterone. Dominance was significantly positively correlated with badge size, but not with other morphological traits. We found little support for the social control hypothesis. Birds did not have significantly more interactions with individuals of similar badge size, before the manipulation. Similarly, after the experimental increase in badge size, experimental birds did not tend to have more encounters with large-badged males. Experimental birds with enlarged badges won more fights compared with prior to the manipulation, suggesting that badge size is used as a signal of social dominance even in small and stable flocks. Finally, corticosterone levels in the blood did not increase significantly after the manipulation of badge size, suggesting that there is no measurable cost, resulting from stress, in cheaters.     

Dominance relations in captive groups of adult and juvenile arctic blue foxes (Alopex lagopus)

Dominance relationships were studied in captive Arctic blue fox (Alopex lagopus) groups comprising adults (four males, five females) and juveniles (four males, five females). The results showed that Arctic blue foxes easily formed a social organization with an observable hierarchy, in which adults typically dominated over juveniles. Within the same age group, males usually dominated over females. Dominance correlated most significantly with body weight in autumn, but later that correlation decreased. Urine marking activity was very low during autumn and early winter, but increased significantly prior to and during the breeding season when aggressive encounters were also most pronounced. In addition to several adults, the social status of some juveniles was high at breeding time. Altogether 7 out of 11 females (63.6%) whelped, but the survival rate of litters was low and kits of only two adult females survived (18.2%). It can be concluded that hierarchical development and reproduction in Arctic blue fox groups are markedly influenced by dominance relationships.     

Social dominance in free-living Willow Tits Parus montanus: determinants and some implications of hierarchy

Several features of social dominance among Willow Tit Parus montanus winter flocks were examined during a four-winter study. Birds of both sexes were evenly distributed over the 33 flocks studied. In nearly half of the flocks there was an adult pair accompanied by yearlings, but one-third of the flocks consisted of more than two adults with yearlings. Males were found to be dominant over and larger than females. Within a sex, yearlings were usually subordinate to adults. The effect of size on dominance, after controlled for sex and age, remained obscure in our field data. The hierarchical status of an individual was found to rise or at least stay the same in different years, which supports the "hopeful dominants" hypothesis, i.e. birds stay in flocks hoping to achieve a higher status in the future. The ranks of mates correlated highly significantly, implying that high-ranking birds were paired with other high-ranking birds and low-ranking birds with other low-ranking birds. Birds of different age and sex did not show any differences in the proportion of initiated aggressive encounters directed at other individuals. However, males were more aggressive to other males than to females and also tended to behave less aggressively towards their own mates than towards other individuals in the flock. This could be a male strategy to strengthen the pair-bond and to enhance mate protection described earlier in     

Plasma androgens and aggressiveness in captive winter white-throated sparrows (Zonotrichia albicollis)

Captive white-throated sparrows (Zonotrichia albicollis), held under a natural photoperiod, were pair tested to evaluate dominance and aggressiveness in relation to plasma androgen levels and plumage morphology. Androgen levels were determined in November, 1 week before behavioral testing began; in January, at the conclusion of behavioral testing; and again in March. November plasma androgen levels of captive birds did not differ from those of a group of feral birds and were greater than androgen levels in January and March. Androgen levels of male and female white-striped and tan-striped birds did not differ for any collection date. November plasma A levels were significantly positively correlated with the display of aggressive behaviors by test birds and were inversely correlated, though not significantly, with hierarchy position. Plasma androgen levels determined in January and March were not related to aggressiveness or dominance. These data suggest that circulating androgens and morphological characters may act synergistically, at the onset of flock formation, to influence avian winter dominance hierarchies.     

Sex-reversed correlation between stress levels and dominance rank in a captive non-breeder flock of crows

Group living has both benefits and costs to individuals; benefits include efficient acquisition of resources, and costs include stress from social conflicts among group members. Such social challenges result in hierarchical dominance ranking among group members as a solution to avoid escalating conflict that causes different levels of basal stress between individuals at different ranks. Stress-associated glucocorticoid (corticosterone in rodents and birds; CORT) levels are known to correlate with dominance rank in diverse taxa and to covary with various social factors, such as sex and dominance maintenance styles. Although there is much evidence for sex differences in the basal levels of CORT in various species, the correlation of sex differences in basal CORT with dominance rank is poorly understood. We investigated the correlation between CORT metabolites (CM) in the droppings and social factors, including rank and sex, in a captive non-breeder group of crows. In this group, all the single males dominated all the single females, and dominance ranks were stable among single males but relatively unstable among single females. CM levels and rank were significantly correlated in a sex-reversed fashion: males at higher rank (i.e., more dominant) had higher CM, whereas females at higher rank exhibited lower CM. This is the first evidence of sex-reversed patterns of CM–rank correlation in birds. The results suggest that different mechanisms of stress–dominance relationships operate on the sexes in non-breeder crow aggregations; in males, stress is associated with the cost of aggressive displays, whereas females experience subordination stress due to males' overt aggression.     

迁地保育林麝的侵犯性、等级序位及相互关系

侵犯性是动物个性的重要维度之一,体现了其主动挑衅和攻击其他个体的倾向。动物的侵犯性与其社会结构及等级序位存在紧密关系。本研究于2018年6月1日至7月31日对四川马尔康林麝繁育场的圈养林麝(Moschus berezovskii)进行了行为取样,计算林麝个体的侵犯性个性(侵犯性指数)和等级序位指数,分析圈养林麝的侵犯性和等级序位格局、影响因素及相互关系。结果表明,圈养林麝的侵犯性在性别间存在显著差异,雄性林麝的侵犯性(0.45±0.09,n=22)显著高于雌性(0.22±0.06,n=30)(P <0.05),年龄和驯养密度对其侵犯性的效应均不显著(P> 0.05),说明林麝侵犯性的刚性较强;圈养林麝等级序位的性别间差异不显著(P> 0.05),亚成体麝与成体麝的等级序位差异也未达显著水平(P> 0.05),原因在于麝场的建群未区分年龄组;圈养林麝个体的侵犯性与其等级序位显著正相关,林麝个体的侵犯性越大,其等级序位越高(r=0.73,P <0.05),推测这与社群的序位等级构建和资源竞争有关。     

Social inertia in white-throated sparrows results from recognition of opponents

Social inertia is a term for the stability of dominance relationships despite changes in the intrinsic dominating abilities of opponents. In a standard test for social inertia, low-ranking birds in an established hierarchy receive implants with testosterone (treated) and high-ranking birds receive empty implants (untreated). Social inertia occurs when the treated birds remain subordinate to untreated opponents in these groups, despite evidence that similarly treated birds dominate untreated strangers. In previous demonstrations of social inertia, however, treated and untreated birds were returned to their original aviaries and tested with familiar opponents, and thus the effects of familiarity with the location and those of familiarity with opponents were not separated. To address this issue, we investigated social inertia in 16 groups of white-throated sparrows Zonotrichia albicollis. When low-ranking treated birds were placed in new aviaries with familiar high-ranking, untreated opponents (treatment S, same opponents), dominance relationships showed social inertia. When such birds were placed in new aviaries with unfamiliar opponents (treatment N, new opponents), testosterone influenced dominance. When groups of high-ranking, untreated birds acquainted with each other were placed with unfamiliar treated opponents (treatment G, grouped dominants), 'coat-tail' effects (dominance by association with high-ranking individuals) sometimes outweighed the effects of testosterone. Social inertia in this species is thus a result of familiarity with opponents, rather than familiarity with locations of encounters. Measurements of aggressive tendencies confirmed a previous report that social inertia suppresses activation of aggression by testosterone. White-throated sparrows can thus recognize their opponents, and this ability affects the expression of both dominance and aggression. Copyright 1999 The Association for the Study of Animal Behaviour.     

Piping as a display of dominance by wintering Oystercatchers Haematopus ostralegus

Although Oystercatchers Haematopus ostralegus wintering in the Exe Estuary performed the piping display in any type of aggressive encounter, it was predominantly observed in two contexts. ‘Solitary piping’ occurred when one individual piped without an obvious opponent being in the immediate vicinity. In ‘piping ceremonies’, two or more individuals piped at each other for prolonged periods at close distance. Prolonged contests over mussels were usually won by the individual that had initially started piping. The most aggressive and successful birds (the dominants), engaged most often in solitary piping and piping ceremonies, using up to 10% of their time this way. This suggests that the behaviour was associated with dominance rank. The hypothesis is that the two birds are likely to start piping if their relative status has not already been established, so that a quick end to a dispute over a mussel or feeding site is not achieved. A dominant individual performs solitary piping from a distance to suppress the aggressive activities of other birds within its feeding range. If this signal fails, the dominant may join the ceremony, explaining why three was the most frequent number of birds involved in piping ceremonies. The ‘butterfly’ flight, used mainly by apparently dominant birds as they arrived on the feeding grounds, may signal their high status to birds already there. The outcome of several piping ceremonies involving birds of known dominance was consistent with this explanation. The location of piping ceremonies involving only two birds (interpreted as piping ceremonies not joined by dominants) at the extremities of the feeding ranges of the most dominant individuals is also consistent with this hypothesis, because this is where their dominance would be expected to decline and would therefore be more likely to be contested. This explanation of piping and butterfly displays assumes that individuals can recognize other individuals or quickly assess their dominance. Since dominant individuals were attacked less often than subdominants and attacking birds were rebuffed less than would be expected if recognition did not occur, we suggest that some form of dominance or individual recognition did exist. The width of the white collar did not correlate with dominance, but the individual variations in this feature may enable birds to distinguish each other and thus learn their relative status. It is intriguing that the diplomatist posture, which is typically used to rebuff misguided attackers, seems almost designed to give the opponent a particularly clear view of the white collar.     

Aggressiveness in king penguins in relation to reproductive status and territory location

King penguins, Aptenodytes patagonicus, vigorously defend small territories in very dense colonies. The egg-laying season lasts approximately 4 months, but only pairs that reproduce during the first half of the period succeed in fledging a chick. I examined various factors affecting aggressiveness of king penguins during the breeding season and focused on the differences between central and peripheral territories. Pairs on peripheral territories experienced twice as many encounters with avian predators as did central birds. The vast majority of peripheral birds were late breeders, indicating that reproductive success was very low among penguins defending a territory on the edge of the colony. Time invested in territory defence and rate of agonistic encounters between breeding neighbours increased from the incubation to the brooding period. Parents gave most threat displays to territorial neighbours when the chick was very young and just before crèche formation. Distance to colony edge was not related to aggressiveness in incubating birds, however, the rate of pecking and flipper blows increased from the edge to the centre during brooding. In addition, aggressiveness of breeding penguins towards travelling birds trespassing into their territory increased with distance to edge. Early breeders were not more aggressive than late breeders but the proportion of time spent in territory defence increased with the number of days a bird had spent incubating. As expected, I did not detect any sex difference in aggressive behaviour. Birds occupying territories on the beach were generally more aggressive during the incubation period than those located on the valley sides. Reproductive status (incubating versus brooding) and territory location were the main factors explaining the various levels of aggressiveness observed in breeding king penguins. Copyright 2000 The Association for the Study of Animal Behaviour.     

WINTER SURVIVAL IN RELATION TO DOMINANCE CLASSES AMONG SILVEREYES ZOSTEROPS LATERALIS CHLOROCEPHALA OF HERON ISLAND, GREAT BARRIER REEF

Winter survival with respect to dominance classes of 932 individually colour-ringed Silvereyes was examined on Heron Island, Great Barrier Reef, between 1965 and 1969. The dominants (winning two-thirds or more of aggressive encounters) had significantly better chances of survival between May and August (southern winter) than other birds. The 1967/68 year group was studied in detail; the young born early in the breeding season contained proportionately more dominants than those born later in the season and dominant birds tended to survive better in winter. Adults in the same period showed no dominance dependent survival. The weight of birds in winter differed between first-year birds and adults in most cases, but winter mortality within each year-group was not related to the weight of individual birds in May. However, the dominant class had a smaller proportion of birds losing weight through the winter than other classes, and the dominant adults and the intermediate class of first-year birds tended to be heavier than others in August. The lengths of wing, tail, tarsus and exposed culmen examined for the 1967/68 year group showed no significant trends in either survival or dominance classes. Better survival of dominant birds is considered to be a consequence of their feeding advantages over others, but the intensity of selection for an ability for dominance may fluctuate from year to year in relation to the population density and distribution and abundance of food supply.     

Karyological study of some Corvine birds (Corvidae, aves)

Karyotypes were studied in the hooded and carrion crows, their naturally occurred hybrids, the jungle crow, the azure-winged magpie (2n = 80 in all aforementioned birds), and the magpie (2n = 82). Corvine birds of Primorskii Krai were karyotyped for the first time. In addition to the similarity in the diploid chromosome sets, corvine birds were shown to have a similar structure of karyotype: in all studied birds, 14 macrochromosomes (Mchs) classified into three groups according to their size were detected. By karyotype structure, birds belonging to the same genus are similar. Some intergeneric differences are due to a change in the position of centromeres of the largest and sex chromosomes. Karyotypes of interspecific hybrids of crows are remarkable for the presence of heteromorphic (t/st) chromosome pair 2 in some individuals, which apparently does not affect their fecundity. Using differential C-banding, the sex chromosome W in female magpies was identified. In addition, heteromorphism was detected in C-bands of homologs of Mch pair 4 in the hooded crow. In the jungle crow, the azure-winged magpie, and the magpie, bright QH-bands and numerous G-bands were detected on Mchs and on some microchromosomes only. Active Ag-NOR-bands were detected on one macrochromosome pair in the magpie. In all, the karyotype structure of corvine birds is comparable to the basic structural scheme of the karyotype in the order Passeriformes, which confirms the concept of conservatism of the avian karyotype.     

The First Description of Dominance Hierarchy in Captive Giraffe: Not Loose and Egalitarian,but Clear and Linear

Wild giraffes live in extensive groups in the fission fusion system, maintaining long social distances and loose social bonds. Within these groups, resources are widely distributed, agonistic encounters are scarce and the dominance hierarchy was reported in males only, while never deeply analysed. In captivity, the possibility to maintain inter-individual distances is limited and part of the resources is not evenly distributed. Consequently, we suggest that agonistic encounters should be more frequent, leading to the establishment of the dominance hierarchy. Based on the differences in resource-holding potential, we suggested that the rank of an individual would be affected by age and sex. Based on hypotheses of prior ownership, we tested whether rank was positively affected by the time spent in a herd and whether it was stable in adult females, which were present long-term in the same herd. We originally monitored four herds of Rothschild giraffes (Giraffa camelopardalis rothschildii) in Dvůr Králové zoo (n = 8), Liberec zoo (n = 6), and two herds in Prague zoo: Prague 1 (n = 8) and Prague 2 (n = 9). The Prague 1 and Prague 2 herds were then combined and the resulting fifth herd was observed over three consecutive years (2009, 2010, and 2011) (n = 14, 13, and 14, respectively). We revealed a significantly linear hierarchy in Dvůr Králové, Prague 2 and in the combined herd in Prague. Rank was significantly affected by age in all herds; older individuals dominated the younger ones. In females, rank was positively affected by the time spent in the herd and adult females in Prague maintained their rank during three consecutive years. This study represents the first analysis of the dominance hierarchy in the captive giraffe, and discusses the behavioural flexibility of the social structure in response to monopolisable resources in a captive environment.     

Linear and Stable Dominance Hierarchies in Cooperative Carrion Crows

Cooperatively breeding birds typically form cohesive and stable groups that live year‐round in all purpose territories where competition for resources is likely to arise. Understanding how group members negotiate over resources is crucial because conflicts may disrupt the stability of the group and may ultimately hinder cooperation. However, social relationships within the group have been largely neglected so far. Here we investigated how cooperatively breeding carrion crows (Corvus corone corone) share a food source, by observing dyadic interactions in 29 territories that contained retained offspring of the breeding pair and/or immigrants. We found that crows formed linear and stable dominance hierarchies, which were stronger for males than females. We suggest that this difference mirrors the level of competition for resources other than food, such as reproduction and territory inheritance, which is higher in males than females. Interestingly, immigrant males dominated male offspring, suggesting that, for the resident breeder, which is the alpha member of the group, the benefits of an association with an immigrant overcome the costs of having his sons pushed down in the hierarchy. Our study uncovered the key factors that determine hierarchical relationships among cooperatively breeding crows and highlighted the need of focussing on social interactions in every context of group living to fully explain the dynamic of cooperation at the nest.     

A Karyological Study of Some Corvine Birds (Corvidae, Aves)

Karyotypes were studied in the hooded and carrion crows, their naturally occurred hybrids, the jungle crow, the azure-winged magpie (2n= 80 in all aforementioned birds), and the magpie (2n= 82). Corvine birds of Primorskii Krai were karyotyped for the first time. In addition to the similarity in the diploid chromosome sets, corvine birds were shown to have a similar structure of karyotype: in all studied birds, 14 macrochromosomes (Mchs) classified into three groups according to their size were detected. By karyotype structure, birds belonging to the same genus are similar. Some intergeneric differences are due to a change in the position of centromeres of the largest and sex chromosomes. Karyotypes of interspecific hybrids of crows are remarkable for the presence of heteromorphic (t/st) chromosome pair 2 in some individuals, which apparently does not affect their fecundity. Using differential C-banding, the sex chromosome W in female magpies was identified. In addition, heteromorphism was detected in C-bands of homologs of Mch pair 4 in the hooded crow. In the jungle crow, the azure-winged magpie, and the magpie, bright QH-bands and numerous G-bands were detected on Mchs and on some microchromosomes only. Active Ag-NOR-bands were detected on one macrochromosome pair in the magpie. In all, the karyotype structure of corvine birds is comparable to the basic structural scheme of the karyotype in the order Passeriformes, which confirms the concept of conservatism of the avian karyotype.     

Social relationships and social structure of the mound-building mouse (<Emphasis Type="Italic">Mus spicilegus</Emphasis>) in intraspecific cage groups

The social relationships of Mus spicilegus were studied in intraspecific cage groups for a month during the reproductive period to clarify the role of intraspecific aggressiveness in shaping social structure and in reproduction. The mound-building mice were captured from a wild population in northern Bulgaria. A total of 13 groups, 5 male, 4 female, and 4 male–female groups, were tested. Each group consisted of five unfamiliar adults. The experiments were carried out in 100×100-cm glass cages. The patterns of behaviour and the dominant–subordinate structure within each group were investigated. A dominance index for every individual in the group was calculated. During the period of establishment of the hierarchical structure in the groups a high level of aggressiveness was registered among individuals. About half of the subdominants and subordinates were found dead, with or without obvious marks of injury. Male–male interactions were more aggressive compared to these of females. Dominant–subordinate relationships between individuals of the same sex were observed in the mixed groups. A link between dominance rank and reproduction was established. The dominant male and the dominant female formed a pair and started to breed. Based on these findings it can be assumed that the aggressiveness of M. spicilegus is an adaptation to its life in the fields, where dominance is likely to be an essential factor in individual reproductive success. It is likely that the successful rearing of offspring in natural populations, which are exposed to strong biotic and abiotic impact, necessitates the existence of monogamous relationships. Communicated by P. Gama Mota     

Categorical learning between 'male' and 'female' photographic human faces in jungle crows (Corvus macrorhynchos)

We trained jungle crows to discriminate among photographs of human face according to their sex in a simultaneous two-alternative task to study their categorical learning ability. Once the crows reached a discrimination criterion (greater than or equal to 80% correct choices in two consecutive sessions; binomial probability test, p < .05), they next received generalization and transfer tests (i.e., greyscale, contour, and ‘full’ occlusion) in Experiment 1 followed by a ‘partial’ occlusion test in Experiment 2 and random stimuli pair test in Experiment 3. Jungle crows learned the discrimination task in a few trials and successfully generalized to novel stimuli sets. However, all crows failed the greyscale test and half of them the contour test. Neither occlusion of internal features of the face, nor randomly pairing of exemplars affected discrimination performance of most, if not all crows. We suggest that jungle crows categorize human face photographs based on perceptual similarities as other non-human animals do, and colour appears to be the most salient feature controlling discriminative behaviour. However, the variability in the use of facial contours among individuals suggests an exploitation of multiple features and individual differences in visual information processing among jungle crows.