Helping at the nest, allofeeding and social status in immature arabian babblers
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Arabian babblers, Turdoides sqamiceps, are cooperatively breeding, group territorial birds, occurring in desert habitats. Non-breeders participate in several types of cooperative behaviour, including care of eggs, nestlings and fledglings, and bringing food to one another (allofeeding). This paper reports on observations of helping with eggs and young, and peer allofeeding, performed by immature babblers, fledged the previous season or earlier in the same season. The birds were from a babbler population studied since 1971 in the Arava valley of eastern Israel. They belonged to groups in which all individuals had been colourringed and were accustomed to human observers. Individuals and nests in the territories of these groups could be watched from distances of 1–2 m without causing alarm, so fine details of behaviour could be observed in the field. In most allofeeding interactions observed, a more dominant individual brought food to a subordinate. Sometimes the subordinate bird avoided accepting the proferred food, and was then hit or chased by the bird that had attempted to feed it. Occasionally a subordinate bird attempted to feed a more dominant individual. Dominant birds always refused to take food offered by a subordinate, and hit or chased any subordinate attempting to feed them. Frequencies of visits to nests were correlated with helper rank for those visits where the incoming bird displaced a previous visiter from the nest. Frequences of visits to unattended nests, however, were not correlated with helper rank. Similarly, frequencies of feeding visits to fledglings were correlated with helper rank until incubation of the next brood began. After this, the correlation disappeared. Numerous instances of aborted nest visits were observed in which a helper, often carrying food, arrived at the nest-tree but left again without visiting the nest. In many other cases, a helper arrived at the nest-tree but delayed visiting the nest for several minutes. Aborted and delayed visits usually occurred when a more dominant bird was in the vicinity of the nest. This suggests interference between helpers. Direct interference between helpers visiting fledglings was occasionally observed. In such cases, a more dominant helper snatched food from a subordinate approaching fledglings, and then fed this food to the fledglings itself. Interference between helpers, and conflict between proferers and recipients of food during allofeeding, are not easily explained by kin selection or reciprocity. On the other hand, such behaviour is readily explained by a hypothesis which suggests that individuals may increase their social status in the group by performing cooperative behaviour. Babblers that establish status by demonstrating ability to bear the short-term costs of cooperative behaviour, rather than through direct aggression towards rivals, are likely to forge collaborative relationships with other group members. Since babblers must collaborate to establish and defend a territory, such relationships are essential to reproductive success.
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