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.     

On the Relationship between Aggression and Social Dominance

It is frequently assumed that variation in dominance status is in large part a function of differences in levels of aggressiveness. This assumption and the reasoning underlying it are examined in this paper. It is argued that the large role ascribed to aggression with respect to dominance success results from a conceptual error, in particular, the temptation to conflate the two constructs. There may be an association between aggression and social dominance, not because levels of aggression determine dominance success, but because aggression is a manifestation of dominance. It is further argued that the analysis of dominance relationships exclusively in terms of individual attributes such as aggression is flawed. Social dominance is a relational quality, which could be profitably studied from a sociological perspective.     

Measuring Dominance and Constructing Hierarchies: An Example Using Mule Deer

Important issues that are still unresolved in the study of animal social groups are how dominance is measured and how individuals are ranked. Based on observations of mule deer (Odocoileus hemionus hemionus), we constructed hierarchies for the same 26 bucks using five potential correlates of dominance and three measures based on the outcome of agonistic interactions. Data for each of six behavior measures were converted to ranks by six different numerical techniques. Different measures of behavior resulted in different hierarchies. This was especially so when the hierarchy based on sparring rates was compared to hierarchies based on other criteria. Although our results showed that dominance is not a unitary concept, several measures of dominance were highly correlated with one another. Thus, one measure, in some instances, may be a good but imperfect predictor of dominance defined by other criteria. For data sets based on index scores or rates of performing behavior, the mean difference and standard-error difference ranking methods had distinct advantages. Both produced equally weighted ranks that minimized the effects of sampling errors.     

Can Individual Differences in History of Dominance Explain the Development of Linear Dominance Hierarchies?

Observed dominance hierarchies are often more linear than expected from randomly-formed dominance relationships, and in triads of animals attacks are distributed non-randomly. I hypothesize that an individual's history of dominance affects its probability of initiating aggressive interactions in the future and that individuals with winning records are more likely to initiate (winning begets initiating). Consistent with this hypothesis, evidence is presented that dominant individuals are more likely to attack than subordinate individuals. The winning begets initiating hypothesis may also explain why correlations between predicted dominance ranks (based on size, age etc.) and observed dominance ranks can be low: If the cost of engaging in and losing an interaction is high relative to the potential benefits of winning, then a large individual conditioned to be subordinate may refrain from contesting smaller, dominant individuals despite its actual competitive superiority.     

Temporal Hierarchies of Variation and the Maintenance of Diversity

Abstract A general model shows how the long-term growth rate of a population can be partitioned into components representing various mechanisms of maintenance of species diversity. One component summarises the effects of fluctuation-independent mechanisms, which include classical resource partitioning and frequency-dependent herbivory. Two other components represent fluctuation-dependent mechanisms, the storage effect and relative nonlinearity of competition.
The general model shows how a community will track an equilibrium set by fluctuation-independent mechanisms and the environmental state when community dynamics are faster than the rate of environmental change. Fluctuation-dependent mechanisms can be important for diversity maintenance with or without such tracking, but on long timescales their effects are indistinguishable from those of fluctuation-independent mechanisms.
These considerations lead to a hierarchical view of mechanisms of diversity maintenance where the effects of different timescales are partitioned or merged depending on the timescale of observation. These issues are illustrated with model examples involving various combinations of resource partitioning, fluctuations in recruitment rates, variation in the timing of germination, and seasonality. The very long timescales associated with climate change contain many complexities but nevertheless many ideas applicable to shorter timescales may be useful in a modified form.     

Modeling Social Dominance: Elo-Ratings,Prior History,and the Intensity of Aggression

Among studies of social species, it is common practice to rank individuals using dyadic social dominance relationships. The Elo-rating method for achieving this is powerful and increasingly popular, particularly among studies of nonhuman primates, but suffers from two deficiencies that hamper its usefulness: an initial burn-in period during which the model is unreliable and an assumption that all win–loss interactions are equivalent in their influence on rank trajectories. Here, I present R code that addresses these deficiencies by incorporating two modifications to a previously published function, testing this with data from a 9-mo observational study of social interactions among wild male chimpanzees (Pan troglodytes) in Uganda. I found that, unmodified, the R function failed to resolve a hierarchy, with the burn-in period spanning much of the study. Using the modified function, I incorporated both prior knowledge of dominance ranks and varying intensities of aggression. This effectively eliminated the burn-in period, generating rank trajectories that were consistent with the direction of pant-grunt vocalizations (an unambiguous demonstration of subordinacy) and field observations, as well as showing a clear relationship between rank and mating success. This function is likely to be particularly useful in studies that are short relative to the frequency of aggressive interactions, for longer-term data sets disrupted by periods of lower quality or missing data, and for projects investigating the relative importance of differing behaviors in driving changes in social dominance. This study highlights the need for caution when using Elo-ratings to model social dominance in nonhuman primates and other species.     

Sex Differences in the Steepness of Dominance Hierarchies in Captive Bonobo Groups

Bonobos have a reputation as a female-dominated and egalitarian species. We examined the 2 aspects of dominance in 6 captive bonobo groups. Females do not consistently evoke submission from all males in all contexts. Though females occupy the highest-ranking positions in the dominance hierarchy, there are in each group males that obtain rather high ranks and are able to dominate ≥1 female. Thus female dominance is not complete and hierarchies can be better described as nonexclusive female dominance. We studied egalitarianism by measuring linearity and steepness of dominance hierarchies. The hierarchies of all groups are highly linear. Hierarchies among males are steeper than among females. On average, male bonobos are more despotic than females, but females too can have despotic relations, both with other females and with males. Hence one can call bonobos in captivity semidespotic rather than egalitarian.     

Mounting Interactions between Female Japanese Macaques: Testing the Influence of Dominance and Aggression

The goal of the work reported here was to determine whether female Japanese macaques (Macaca fuscata) participated in same-sex mounting interactions during homosexual consortships to communicate about asymmetries in their dominance relationships and to reduce aggression. Focal data were collected during 21 homosexual consortships involving 14 females living in a captive, mixed-sex group of 37 individuals. We identified eight types of mounts, one solicitation used specifically to request to mount (hands-on-hindquarters solicitation), two solicitations used specifically to request to be mounted (hindquarter and back presentations), and one behavior employed to facilitate mounts-in-progress (clasping). We tested whether dominant consort partners (1) mounted more and (2) requested to mount more than their subordinate partners and whether subordinate consort partners (1) requested to be mounted more and (2) facilitated mounts-in-progress more than their dominant partners. Finally, we examined whether mounting was temporally linked to the onset of aggressive interactions between consort partners and whether it functioned to defuse incipient aggression. None of these predictions was supported. All types of mounts, mount solicitations, and clasping occurred bi-directionally within consort dyads. Mutual sexual attraction and gratification provided the proximate motivation for these mounting interactions and, in turn, for the formation and maintenance of their homosexual consortships.     

Large size in an island-dwelling bird: intraspecific competition and the Dominance Hypothesis

Differences between island‐ and mainland‐dwelling forms provide several classic ecological puzzles. Why, for instance, are island‐dwelling passerine birds consistently larger than their mainland counterparts? We examine the ‘Dominance hypothesis’, based on intraspecific competition, which states that large size in island passerines evolves through selection for success in agonistic encounters. We use the Heron Island population of Capricorn silvereyes (Zosterops lateralis chlorocephalus), a large‐bodied island‐dwelling race of white‐eye (Zosteropidae), to test three assumptions of this hypothesis; that (i) large size is positively associated with high fitness, (ii) large size is associated with dominance, and (iii) the relationship between size and dominance is particularly pronounced under extreme intraspecific competition. Our results supported the first two of these assumptions, but provided mixed evidence on the third. On balance, we suggest that the Dominance Hypothesis is a plausible mechanism for the evolution of large size of island passerines, but urge further empirical tests on the role of intraspecific competition on oceanic islands versus that on mainlands.     

Social Behaviour in a Colony of Domestic Guinea Pigs: Aggression and Dominance

Social behaviour, particularly dominance behaviour, in a colony of domestic guinea pigs was investigated. Observations were of four different types, (1) daily observations on the whole group; (2) interactions between pairs of ♂♂ (in all possible combinations) were recorded when competing over a single water source; (3) interactions between all the ♂♂ in the group observed together at a single water source; (4) as (2) but in the absence of water, in order to observe fighting. It was hoped that the study would demonstrate the extent to which a hierarchial structure, if one such existed, was consistent in different social contexts.     

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.     

Hormonal Correlates of Dominance and Starvation-induced Aggression in Chicks of the Blue-footed Booby

In the blue-footed booby (Sula nebouxii), the first-hatched chick aggressively dominates its sibling and sometimes kills it when food is in short supply. To investigate the endocrine correlates of dominance-subordinance and hunger-induced agonism, we deprived 15–20-d-old single-chick and two-chick broods of food during 48 h by taping chicks' necks to prevent ingestion of parentally provided food (a protocol used previously and known to elicit escalated sibling fighting). We monitored weight and levels of circulating testosterone, oestradiol and corticosterone in deprived and normally fed broods comprising singletons, seniors (= dominants) and juniors (= subordinates), and observed behaviour to verify that aggression increased in deprived two-chick broods. During the 2 d of fasting, experimental chicks lost on average 6% of their baseline weight. After normal feeding was reinstated, seniors and singletons recovered normal (control) weight, but juniors remained significantly lighter than controls. No testosterone was detected in any nestling, but baseline corticosterone level was 109% higher in juniors than in seniors or singletons, implying that elevated corticosterone in juniors is a consequence of social subordination and may facilitate submissive behaviour. Although there was evidence that aggression of seniors increased under food deprivation, the increase was not accompanied by the rise in levels of testosterone expected under the ‘Challenge hypothesis’ (Wingfield et al. 1990). This result implies that this hypothesis probably does not apply to booby nestlings in the context of starvation-induced aggression. During the 2 d deprivation period, corticosterone levels of experimental chicks increased significantly, then declined to baseline levels a day after tapes were removed. The increase was probably due to the combined effects of starvation and frustration. We suggest that corticosterone could alter responsiveness of nestling blue-footed boobies to external stimuli, resulting in more aggression by dominants and greater readiness of subordinates to submit.     

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.     

FBD Marriage: Further Support for the Westermarck Hypothesis of the Incest Taboo?

In the past decade, several reports, including data from Taiwan on the sim-pua form of marriage, have provided evidence for the Westermarck Hypothesis, a heretofore unpopular explanation of the incest taboo. This theory states that intimate childhood association breeds sexual disinterest. A preliminary study of FBD marriage in Lebanon reveals similar support for Westermarck's theory. The Lebanese patrilateral parallel cousin marriages examined produced significantly fewer children and more divorces than nonpaternal first cousin unions. Given the quite disparate cultures from which this evidence comes, serious reconsideration of Westermarck's Hypothesis is suggested. [incest taboo, FBD marriage, Lebanon, Middle Eastern sexuality]     

Dominance,Age and Aggression among Female Pronghorn,Antilocapra americana (Family: Antilocapridae)

The relationships among dominance, age and aggressive behaviour of marked, female pronghorn (Antilocapra americana; Family Antilocapridae) were studied in north central Colorado. Females formed dominance hierarchies with few circular relationships. Unlike other female ungulates, dominance rank was not correlated with age. Dominance rank was negatively correlated with rate of aggression initiated in summer and winter; however, aggressiveness as an individual trait was not related to dominance. Older females received higher rates of aggression than younger females in summer and winter. During the rut, dominance rank was correlated with rate of aggression received but not with rate of aggression initiated. Patterns of aggression that differ by season within a given hierarchy of dominance relationships suggest a different cause or function of aggressive behaviour in different seasons.     

Maternal Investment in the Swordtail Fish Xiphophorus multilineatus: Support for the Differential Allocation Hypothesis

The differential allocation hypothesis predicts that reproductive investment will be influenced by mate attractiveness, given a cost to reproduction and a tradeoff between current and future reproduction. We tested the differential allocation hypothesis in the swordtail fish Xiphophorus multilineatus, where males have genetically influenced (patroclinous inheritance) alternative mating tactics (ARTs) maintained by a tradeoff between being more attractive to females (mature later as larger courting males) and a higher probability of reaching sexual maturity (mature earlier as smaller sneaker males). Males in X. multilineatus do not provide parental care or other resources to the offspring. Allelic variation and copy number of the Mc4R gene on the Y-chromosome influences the size differences between males, however there is no variation in this gene on the X-chromosome. Therefore, to determine if mothers invested more in offspring of the larger courter males, we examined age to sexual maturity for daughters. We confirmed a tradeoff between number of offspring and female offspring’s age to sexual maturity, corroborating that there is a cost to reproduction. In addition, the ART of their fathers significantly influenced the age at which daughters reached sexual maturity, suggesting increased maternal investment to daughters of courter males. The differential allocation we detected was influenced by how long the wild-caught mother had been in the laboratory, as there was a brood order by father genotype (ART) interaction. These results suggest that females can adjust their reproductive investment strategy, and that differential allocation is context specific. We hypothesize that one of two aspects of laboratory conditions produced this shift: increased female condition due to higher quality diet, and/or assessment of future mating opportunities due to isolation from males.     

A Simple and Stable Silica Gel Support for the Oligodeoxyribonucleotide Synthesis

Abstract

A simple and efficient synthesis of a solid support with a long chain polyamide spacer has been developed. The spacer was made by successive addition of ethylene-diamine and succinic anhydride. The obtained solid support gives very homogeneous 20 mer oligodeoxyribonucleotides, detected by HPLC and electrophoresis.     

A Novel Method of Assessing Dominance Hierarchies Shows Nuance,Linearity and Stability in the Dinosaur Ant Dinoponera quadriceps

Many social species with relatively simple societies have dominance hierarchies of individuals, with dominant individuals achieving fitness and subordinate individuals either queuing to obtain fitness or achieving only indirect fitness by helping relatives. Assessing the dominance hierarchy in a social group is generally based upon observing dyadic interactions as and when they occur spontaneously within the whole‐group setting. However, this method can be very time‐consuming because many dyads interact only very rarely, necessitating either extremely long observation periods or many dyadic relationships being unresolved. Here, we report an alternative method using the queenless dinosaur ant Dinoponera quadriceps, which lives in colonies containing tens of individuals. We removed all individuals from their nest and observed the dominance behaviours expressed in isolated dyadic interactions for every pairwise combination of individuals. Individuals showed a classic dominance behaviour in this setting, and the rapid nature of the assay allowed us to observe every dyadic relationship on a weekly basis over 4 weeks. The dominance hierarchies based on these isolated dyadic interactions correlated well with those produced by the conventional method of colony observations. They showed the hierarchies to be highly linear and stable, and also revealed that dominance relationships may extend further down the hierarchy than previously thought. Although highly manipulative, the isolated dyadic interaction method works well and will likely make more feasible the study of other social species in which pairs of individuals can be isolated together.