Fitness increases with partner and neighbour allopreening

Altruism and selfishness are fundamental characteristics of human and animal societies. Among colonial biparental species, breeding outcome depends on interactions between mates and neighbours. However, the relationships between cooperation within and among partnerships and fitness have not been fully investigated. We show that in the highly colonial common guillemot (Uria aalge), altruistic behaviour (allopreening) towards a mate was positively related to long-term fitness, whereas allopreening a neighbour was related to current fitness. Turnover is much lower within than between pairs, so our results suggest that allopreening within pairs generates fitness returns at longer timescales than between pairs. Allopreening not only removes ectoparasites and maintains plumage condition, but may also have important social functions. We found a negative relationship between fight rate and allopreen rate between breeding neighbours, with nests exhibiting low breeding success having a higher frequency of fights with neighbours. We also found evidence for reciprocity in allopreening. Thus, allopreening may function as a reciprocal stress reducer, to decrease the likelihood of fights and associated breeding failure. We suggest that altruistic behaviour has long-term benefits for the survival of the offspring when living in a crowded neighbourhood.     

Social interactions among breeding Brünnich's Guillemots Uria lomvia suggest constraints in relation to offspring vulnerability

Brünnich's Guillemots Uria lomvia are adapted to high-density breeding in large colonies on steep cliffs. Because they breed on narrow ledges, egg loss through dislodgement is an important cause of breeding failure. Fighting among breeders presumably raises the risk of accidental dislodgement. In this study, we investigated whether social behaviour among Brünnich's Guillemots shows any adaptations to reduce accidental egg loss by modifying behaviour during incubation. We found that the amount of aggression increased significantly at the time of hatching, perhaps in response to the reduced risk of breeding failure through dislodgement of a chick, compared with an egg. Allopreening followed an inverse trend, falling significantly after the day of hatch. This supports the hypothesis that allopreening is used to reduce aggressive interactions. At the same time, the frequency of allopreening was greatly increased on days when mosquitoes affected the birds, consistent with the hypothesis that allopreening is part of a defence strategy against ectoparasites.     

ON THE ROLE OF ALLOPREENING INVITATION BEHAVIOUR IN REDUCING AGGRESSION AMONG RED AVADAVATS, WITH COMMENTS ON ITS EVOLUTION IN THE SPERMESTIDAE

This paper discusses the occurence of allopreening and allopreening invitation behaviour in red avadavats, and analyses tho factors which cause these birds to groom partners. It was found that allopreening tended to be temporally linked with aggression and autopreening beaviour in the actor, but that on many occasions, no signs of aggression preceded the act of allopreening. Allopreening invitation behaviour with the head feathers fully raised occurred in situations when the inviting individual was attacked and in response to being pecked or allopreened.
Alloprcening bouts which are tomporally correlated with autopreening behaviour may be a response by the groomer to the feathers of the clumping partner whon the preening tendency of the former bird is alroady activated.
From tho evidence submitted, it seems as though the allopreening invitation posture elicits non-aggressive preening in a bird which is likely to attack. It thus appears to act as an appeasement gesture; head ruffling changes the motivation of a bird tending to be aggressive. It was also found, however, that allopreening invitation behaviour may occur independently of appeasement.
The evolution of the Spermestid allopreening invitation posture from a "fluffed" body posture is also discussed.     

Duration and outcome of intergroup conflict influences intragroup affiliative behaviour

Theoreticians have long suggested that the amount of intergroup conflict in which a group is involved could influence the level of cooperation or affiliation displayed by its members. Despite the prevalence of intergroup conflicts in many social animal species, however, few empirical studies have investigated this potential link. Here, I show that intragroup allopreening rates are highest in green woodhoopoe (Phoeniculus purpureus) groups that have the greatest involvement in intergroup conflict. One reason for this relationship is a post-conflict increase in allopreening, and I demonstrate for the first time that both conflict duration and outcome influence subsequent allopreening rates: group members allopreened more following long conflicts and those they lost compared with short conflicts and those they won, perhaps because the former are more stressful. The increase in affiliative behaviour was the result of more allopreening of subordinate helpers by the dominant breeding pair, which may be because the breeders are trying to encourage helpers to participate in future conflicts; relative group size influences conflict outcome and helpers contribute more to conflicts than do the breeding pair. These results emphasize that our understanding of cooperation and group dynamics can be enhanced by investigations of how intergroup interactions affect intragroup processes.     

Bowing and allopreening of captive Jackdaws Corvus monedula

Allopreening interactions were observed in groups of captive, hand-reared Jackdaws Corvus monedula. Head fluff and Bowing, both resembling 'soliciting' postures described in the literature, are described. Head fluff seems to be associated with allopreening. Bowing was performed more frequently by socially top-ranking birds, performed and received equally frequently by females and by males, and directed at birds higher than and lower than the performer with equal frequency. Bowing was followed mostly by the recipient's moving away or by no apparent response, and only rarely by the allopreening of the actor. Allopreening was performed mostly by socially low-ranking towards high-ranking birds, and both when the actor was dominant to or submissive to the recipient. Bowing and allopreening seemed to be associated with situations of tension. It is suggested that Bowing here was used as intimidation rather than 'solicitation' or appeasement, and that allopreening and Bowing function in confirming relationships and in conveying information about relative status to other members of the group.     

Preparing for battle? Potential intergroup conflict promotes current intragroup affiliation

Groups of human soldiers increase their affiliative behaviour when moving into combat zones. Despite numerous other species also competing as groups, little is known about how potential intergroup conflict might influence current intragroup affiliative behaviour in non-human animals. Here, I show that allopreening (when one individual preens another) increases in groups of cooperatively breeding green woodhoopoes (Phoeniculus purpureus) when they enter areas where conflicts with neighbours are more likely. Self-preening, which is an indicator of stress in other species, did not increase in conflict areas, suggesting that the change in affiliative behaviour is not the simple consequence of greater stress. Instead, because it is the dominant breeding pair that increase their preening of subordinate helpers, it is possible that current affiliative behaviour is being exchanged for agonistic support in any intergroup conflicts that might ensue. These results are important for our understanding of group dynamics, cooperation and the evolution of sociality, but also bring to mind the intriguing possibilities of social contracts and future planning in birds.     

Gender effects on aggression, dominance rank, and affiliative behaviors in a flock of captive adult cockatiels (Nymphicus hollandicus)

Agonistic and affiliative behaviors and spatial positioning were studied in a small psittacine species, Nymphicus hollandicus. Subjects studied were flock-housed breeders including five hens and seven cocks. Fifteen-minute focal animal samples were collected for the entire flock in a randomly distributed order during mate selection and the onset of the breeding season. All agonistic behaviors were recorded, including the winner and loser of each interaction, along with allopreening and copulation behavior. Point samples were recorded every 60s to determine social spacing. Rates of aggression were significantly higher for male cockatiels than female cockatiels. Results based on dyadic agonistic interactions showed males to rank significantly higher in the social hierarchy than females. Associations within the flock were not random. Individual birds associated more with specific birds than would be predicted by chance. Analysis of spatial data revealed that there were both same-sex and opposite-sex preferred associations within the flock. Male cockatiels engaged in allopreening behavior with females significantly more than with other males. Observations of breeding and nesting behaviors revealed pairs, a triad, and extra-pair mating.     

Pair and group breeding behaviour of a communal gallinule,the pukeko, Porphyrio P. Melanotus

Pukeko, Porphyrio porphyrio melanotus breed as pairs or groups and the breeding behaviour of individuals in all units is described and compared. Courtship, including copulation, occurred between all adult members in a territory although male-female copulations were most frequent. Low ranking yearlings were excluded from copulation. Participation in copulation by more than two birds was common, and, along with homosexual behaviour, is believed to help synchronize sexual cycles allowing several females to lay in the same nest at the same time. Only adults incubated, while all birds in a territory assisted with chick care, the degree of participation varying with status. In pair territories, older chicks helped feed younger chicks. Stable kin groups were more coordinated and showed less competitive aggression among members than non-kin groups of unstable membership. The significance of these results is discussed and it is argued that in a situation where an adult group is necessary for the defence of a breeding territory, then it is in the interests of all these individuals to participate in all breeding activities.     

Experimental evidence that sentinel behaviour is affected by risk

Sentinels are a conspicuous feature of some cooperative societies and are often assumed to provide benefits in terms of increased predator detection. Similar to other cooperative behaviours, variation in investment in sentinel behaviour should reflect variation in the benefits of such behaviour. However, evidence for this is inconclusive: to date experiments have manipulated the cost of sentinel behaviour, and considerations of changes in the benefits of sentinel activity on investment patterns are lacking. Here, we experimentally manipulated the benefits of sentinel behaviour in the cooperatively breeding pied babbler (Turdoides bicolor) to assess whether this had any impact on sentinel activity. We simulated the presence of an unseen predator using playbacks of heterospecific alarm calls, and the presence of an actual predator using a model snake. In both cases, the increase in perceived predation risk caused an increase in sentinel activity, demonstrating that investment in sentinel activity increases when the benefits are greater.     

Stress hormones in relation to breeding status and territory location in colonial king penguin: a role for social density?

Because glucocorticoid (stress) hormones fundamentally affect various aspects of the behaviour, life history and fitness of free-living vertebrates, there is a need to understand the environmental factors shaping their variation in natural populations. Here, we examined whether spatial heterogeneity in breeding territory quality affected the stress of colonial king penguin (Aptenodytes patagonicus). We assessed the effects of local climate (wind, sun and ambient temperature) and social conditions (number of neighbours, distance to neighbours) on the baseline levels of plasma total corticosterone (CORT) in 77 incubating and 42 chick-brooding birds, breeding on territories of central or peripheral colony location. We also assessed the oxidative stress status of a sub-sample of central vs. peripheral chick-brooders to determine whether chronic stress arose from breeding on specific territories. On average, we found that brooders had 55 % higher CORT levels than incubators. Regardless of breeding status, central birds experienced greater social density (higher number of neighbours, shorter distance between territories) and had higher CORT levels than peripheral birds. Increasing social density positively explained 40 % of the variation in CORT levels of both incubators and brooders, but the effect was more pronounced in brooders. In contrast, climate was similar among breeding territories and did not significantly affect the CORT levels of breeding birds. In brooders, oxidative stress status was not affected by local density or weather conditions. These results highlight that local heterogeneity in breeding (including social) conditions may strongly affect the stress levels of breeding seabirds. The fitness consequences of such variation remain to be investigated.     

Different patterns of allopreening in the same-sex and opposite-sex interactions of juvenile large-billed crows (Corvus macrorhynchos)

Allogrooming, where an individual grooms another, has been extensively studied in various social animals to understand its role in the evolution of cooperation/prosociality. In existing studies in mammals, allogrooming has been suggested to exhibit not only a hygiene but also a social function. Allopreening, a topic of increasing interest in mammals but recently also in birds, has been studied mostly with mature animals. However, in some species immature individuals also show allopreening and its function remains poorly understood. Crows, Corvus spp., are an ideal model to study this phenomenon, because juveniles form year-round aggregates during their long juvenile stage (e.g., throughout 3–4 years). Here, we investigated the function of allopreening in juvenile groups of wild-caught large-billed crows (C. macrorhynchos). Allopreening frequency and duration for three groups of wild-caught juveniles were analysed to determine whether there was a symmetrical (i.e., reciprocal) or asymmetrical allopreening pattern, and if sex composition of the dyad and/or relative dominance of donor and recipient had an effect. We found that both the frequency and duration of male allopreening correlated with frequency of aggression. Allopreening between both males and females occurred unidirectionally from dominants to subordinates but not in the opposite direction. On the contrary, allopreening between a male and a female was found to be reciprocated, though the absolute frequency and duration were both greater in males than in females. These results suggest that the social function of allopreening in juvenile crows differs depending on the sex composition of the dyad, functioning as a dominance signal for same-sex dyads, and serving a social bonding function for opposite-sex dyads. These findings may reflect the potentially crucial roles of allopreening in within-sex competition and opposite-sex attraction during the 3 year-long juvenile stage affecting future mate choice in lifelong monogamy.     

Sperm traits in relation to male quality in colonial spawning bluegill

Sperm traits (morphology, motility and concentration within ejaculates) and various correlates of male quality (age, body condition, spawning location and timing) were studied in bluegill Lepomis macrochirus , breeding in both the interior and periphery of six colonies in Lake Opinicon, Ontario, Canada. Sperm traits varied significantly more among than within males suggesting that some aspect of male phenotype might influence sperm morphology and behaviour. No measures of male body condition or size were correlated with any sperm or ejaculate traits, when controlling statistically for confounding variables. Sperm swimming speed increased significantly with male age and varied significantly among spawning bouts (controlling for sperm tail length) suggesting that some unknown aspects of male quality might influence the fertilization capacity of spermatozoa. Sperm concentration in ejaculates was significantly higher in males nesting in the interior rather than the periphery of a colony suggesting that those males might also have higher fertilization capacity correlated with their superior dominance status or the lower risk of sperm competition. Thus, older males nesting in the interior of a colony during the first spawning bout of the season are expected to be the best sperm competitors in this population, but the physiological reasons for this increased fertilization capacity remain unknown.     

The relative influence of age and weight on the reproductive behaviour of male grey seals Halichoerus grypus

The breeding behaviour of male grey seals Halichoerus grypus of known age and weight was studied on Sable Island, Nova Scotia. Branded males ranged in age from 8 to 16 years and although there was a positive relationship between age and weight, there was a large overlap between age classes. In general, there was an increase in length of tenure and rates of copulation with increasing male age but this was less marked among males between the ages of 13 and 16 years. Compared to older ones, males of 8 and 12 years were observed at more sites, travelled more extensively around the breeding colony, were transient more frequently and were unable to lie as close to females. There was no correlation between male body weight and either length of tenure or rates of copulation when males of age 8 (the smallest males) were excluded from the analysis. Most agonistic behaviour was made by larger, older males towards smaller, younger ones and appeared to be responsible for the short length of tenure and low reproductive rates of young bulls.     

Sex, Rank and Age Differences in the Japanese Macaque (Macaca fuscata yakui) Participation in Inter-Group Encounters

In many species interactions among group are often characterized by agonistic behaviour. Although animals may participate in inter‐group encounters in different ways, depending on their energetic requirements, reproductive tactics, and/or developmental stage, the proximate causes affecting an animal's participation in inter‐group encounters are still poorly understood. Indeed, many studies have analysed the behaviour of males and females during inter‐group encounters without considering the importance of additional factors (e.g. rank). This study focuses on wild non‐provisioned Japanese macaques (Macaca fuscata yakui) living on Yakushima Island, Japan. It aims to determine how monkeys of different sex, age, and rank behave during inter‐group encounters and it discusses the implications and consequences of their behaviour on group composition and male dispersal. Males participated significantly more than females in inter‐group encounters, by displaying more aggressive or affiliative behaviour. High‐ranking and/or adult males were more aggressive than low‐ranking and/or subadult males during encounters occurring in the mating season and they also showed more herding behaviour. This trend was not found in inter‐group encounters occurring during the non‐mating season. Finally, males which then emigrated to new groups were low‐ranking and/or subadult individuals. Those males displayed more affiliative behaviour towards foreign males than males which did emigrate. These data indicate that in non‐territorial species with male dominance over female and high competition for mating partners males play an active, and often aggressive, role during inter‐group encounter while female participation is scarce. Factors such as age, rank and period of the year (in seasonally breeding species) have to be taken into considerations when analysing interactions between groups and their effects on group composition and social behaviour.     

Die Sozialisation beim Rosenk?pfchenAgapornis roseicollis

Zusammenfassung Junge Rosenköpfchen,Agapornis roseicollis, verlassen durchschnittlich mit 42 Lebenstagen zum erstenmal die Nisthöhlen und werden danach noch etwa zwei Wochen vorwiegend vom Vater gefüttert. Fütterungen treten gelegentlich auch zwischen Geschwistern auf. Die Jungvögel haben mit den Eltern und Geschwistern bis Anfang des vierten Lebensmonats Kraulkontakte. Noch im Jugendkleid, zumeist im Alter von 65 bis 90 Tagen, verpaaren sich Rosenköpfchen mit einem anderen Jungvogel, bevorzugt aus einer anderen Familie und anderen Geschlechts. Diese Jugendpaare bleiben stabil bis über die Geschlechtsreife hinaus. Bestimmte soziale Verhaltensweisen (Sexualverhalten, Kraulen, Kontaktsitzen) werden nur mit dem Partner ausgeführt.

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Socialization of the Peach-faced Lovebird,Agapornis roseicollis

Summary Young Peach-faced Lovebirds, leave their nest-box on an average with 42 days of age. They are still fed for about another two weeks, mainly by the father. Feeding interactions sometimes also occur between siblings. The young undergo allopreening interactions with parents and siblings until the beginning of the fourth month of life. While still in juvenile plumage, usually between 65 und 90 days of life, they form a pair-bond with another young bird, preferably from another family and the other sex. These pairs remain stable beyond sexual maturation. Several social behaviour patterns (sexual behaviour, allopreening, contact sitting) are restricted to the mate.

     

乳腺导管原位癌行前哨淋巴结活检指征的探讨

目的:分析乳腺导管原位癌的前哨淋巴结临床及病理特征,探讨乳腺导管原位癌患者实施前哨淋巴结活检的指征。方法:回顾性分析2002年10月到2010年11月期间诊断为乳腺导管原位癌并行前哨淋巴结活检的46例患者的的前哨淋巴结状态及其与其他临床及病理特征的关系。结果:患者的年龄、肿瘤大小、检出SLN数、肿瘤分级、切缘状态、DCIS类型、是否伴随坏死均无显著统计学意义(P0.05)。可能与乳腺导管原位癌患者前哨淋巴结阳性的几个因素为:年龄41岁-69岁、肿瘤大小1.1 cm-5.0cm、切缘状态未知、肿瘤坏死伴随或未知、肿瘤分级Ⅰ级以上或未知。结论:以下几个因素可能对预测DCIS患者可能出现SLN阳性提供帮助,临床在遇到出现这些因素的DCIS患者需慎重考虑其治疗策略,这些因素包括:年龄41岁-69岁、肿瘤大小1.1cm-5.0 cm、切缘状态未知、肿瘤坏死伴随或未知、肿瘤分级Ⅰ级以上或未知。     

Hormonal and behavioural correlates of male dominance and reproductive status in captive colonies of the naked mole-rat,Heterocephalus glaber.

Naked mole-rat colonies are societies with a high reproductive skew, breeding being restricted to one dominant female (the ''queen'') and 1-3 males. Other colony members of both sexes are reproductively suppressed. Experimental removal of breeding males allowed us to investigate the relationship between urinary testosterone and cortisol, dominance rank, and male reproductive status. Dominance rank was strongly correlated with body weight, age, and urinary testosterone titres in males. No relationship between urinary cortisol levels and male reproductive status or dominance was found. Breeding males were among the highest-ranking, heaviest and oldest males in their respective colonies, and were succeeded by other high-ranking, large, old colony males. In contrast to females, no evidence of competition over breeding status was observed among males. Male-male agonism was low both before and after removal of breeders and mate guarding was not observed. The lower reproductive skew for males compared with female skew or queen control over male reproduction may explain why males compete less strongly than females over breeding status after removal of same-sexed breeders.     

Social relationships and individual contribution to cooperative behaviour in captive common marmosets (Callithrix jacchus)

The social relationships between the members of a family consisting of eight captive common marmosets (Callithrix jacchus) are here examined, as well as the participation of individual family members in infant carrying and defensive behaviour against non-related intruders (adolescent males). Within the social relationships, a clear division of the family into three subgroups (parents, adult offspring, non-adult offspring) could be determined, whereby the adult offspring, especially with respect to the parents, occupy a peripheral position. In the types of cooperative behaviour examined here, substantial participation in infant carrying could be observed in only four family members (parents, adult son, one subadult daughter), and in defence against intruders in only two family members (adult son, one subadult daughter). Participation in infant carrying remained stable throughout the study period. Conversely, individual changes in participation in defence against intruders could be determined, dependent upon the presence of infants carried. The advantages and disadvantages of social substructuring observed are discussed here, in particular the peripheral position of adult offspring, possible connections in individually varied participation in cooperative behaviour, and possible regulative mechanisms. In view of past studies on cooperative behaviour among marmosets, it is here presumed that pronounced individual differences result from a division of labour within the family with respect to various aspects of cooperative behaviour.     

Breeding and management of the common trumpeter (Psophia crepitans)

The breeding biology and management of three wild-caught adult Common trumpeters (one male and two females) was documented at the Woodland Park Zoological Gardens, Seattle, from April, 1984 to August, 1986. A total of 27 eggs were laid, and eight young survived to fledging. Both sexes exhibited crane-like dances during courtship, but the male appeared to perform these behaviors more frequently and with greater intensity than the females. Courtship feeding and allo-preening also occurred. Nests consisted of simple scrapes on the ground, but the birds had no opportunity to rest in trees or other elevated sites. Clutches contained two or three eggs, and the incubation period was aproximately 28 days. Although the breeding pair was generally aggressive toward the second female, all three adults participated in incubation and in caring for the young. Parental behavior consisted of brooding, allopreening, and feeding. The male preened and fed one chick significantly more often than either of the females. Trumpeter chicks were highly precocial, but grew relatively slowly, reaching 50% of adult weight by 45 to 50 days of age. Trumpeters are difficult to maintain and breed in captivity and appear to be susceptible to mycotic diseases, such as aspergillosis. Changes in the social composition of captive groups may result in improved breeding.     

Female-mediated causes and consequences of status change in a social fish

In highly social species, dominant individuals often monopolize reproduction, resulting in reproductive investment that is status dependent. Yet, for subordinates, who typically invest less in reproduction, social status can change and opportunities to ascend to dominant social positions are presented suddenly, requiring abrupt changes in behaviour and physiology. In this study, we examined male reproductive anatomy, physiology and behaviour following experimental manipulations of social status in the cooperatively breeding cichlid fish, Neolamprologus pulcher. This unusual fish species lives in permanent social groups composed of a dominant breeding pair and 1-20 subordinates that form a linear social dominance hierarchy. By removing male breeders, we created 18 breeding vacancies and thus provided an opportunity for subordinate males to ascend in status. Dominant females play an important role in regulating status change, as males successfully ascended to breeder status only when they were slightly larger than the female breeder in their social group. Ascending males rapidly assumed behavioural dominance, demonstrated elevated gonadal investment and androgen concentrations compared with males remaining socially subordinate. Interestingly, to increase gonadal investment ascending males appeared to temporarily restrain somatic growth. These results highlight the complex interactions between social status, reproductive physiology and group dynamics, and underscore a convergent pattern of reproductive investment among highly social, cooperative species.