Review Paper: Daughters on request: about helpers and egg sexes in the Seychelles warbler

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Abstract
The Seychelles warbler (Acrocephalus sechellensis) was an endangered endemic of the Seychelles islands where, until 1988, the entire population of ca. 320 birds was restricted to the one island of Cousin Island (29 ha). Additional breeding populations were successfully established on the islands of Aride (68 ha, 1988) and Cousine (26 ha, 1990) and now with the existence of ca. 2000 warblers on three islands the conservation status of the warbler has improved from endangered to vulnerable. Emigration from the island is extremely rare, so birds that disappeared were known to have died. Almost every bird on Cousin Island has been individually colour ringed and monitored throughout all breeding attempts during a 17–year period (1985–2002; total ca. 2400 birds). These birds were also blood sampled for molecular parentage and sex analyses. Therefore the lifetime reproductive success of many birds is known. Although warblers can breed independently in their first year, some individuals remain in their natal territory as subordinates, and often help by providing nourishment to non–descendant offspring. The frequency of ‘helping’ is affected by habitat saturation and variation in territory quality (insect prey availability). The long–term benefits of helping are higher for daughters than for sons, and it is therefore no wonder that most helpers are daughters from previous broods. Furthermore, on low–quality territories breeding pairs raising sons gain higher fitness benefits than by raising daughters, and vice versa on high–quality territories. Female breeders adaptively modify the sex of their single–egg clutches according to territory quality: male eggs on low quality and female eggs on high quality. The Seychelles warbler is a beautiful example of behavioural and life–history adaptations to restricted circumstances.