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All nests were collected from first broods.
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Three days later adults were removed from second brood bottles and discarded, and purine was added to these bottles.
Two or three broods may be raised each year; birds breeding in colonies produce more eggs and fledglings from their first broods than solitary pairs, but the reverse is true for second and third clutches.
Occasionally, first-year birds from the first brood will assist in feeding the second brood.
Occasionally, young birds from the first brood will assist in feeding the second brood.
There are normally two broods, except in Siberia, where there is only one, the male feeding the fledglings from the first brood while the female sits on the second clutch.
Following Duchateau et al. [ 5], diploid male colonies were identified as those producing workers and males in approximately equal numbers from the first brood.
As diploid males are produced from the first brood, they will be found on the wing, even if the colony from which they have been produced dies prematurely, as our results suggest is highly likely.
These comparisons were restricted to first broods only since paternity data from 2003 was only available for first broods.
We only manipulated first broods, second broods and replacement broods of known first broods after failures were left unmanipulated.
Paternity was assigned to all young in 53 first broods and 4 renestings in 2003, and to all young in 58 first broods and 24 renestings or second broods in 2004.
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