Abstract
Rainfall seasonality is likely an important cue for timing key annual cycle events like moult in birds living in seasonally arid environments, but its precise effect is difficult to establish because seasonal rainfall may affect other covarying annual events such as breeding in the same way. In central Nigeria, however, Common Bulbuls Pycnonotus barbatus moult in the wet season but only show weak breeding seasonality. This suggests that moult is more sensitive to rainfall than breeding, but a similar outcome is possible if moult is simply periodic. We tested the relationship between rainfall and moult in Common Bulbuls at a single location over 18 years: on average moult started 5th May (± 41 days: 25th March–15th June), being on average later than the onset of the rains which is usually mid-April. The likelihood of finding a moulting Common bulbul was best predicted by rainfall 9–15 weeks before moult was scored. We then tested the generality of this across populations: the progress of moult should, therefore, correlate with the average timing of the wet season along a spatial environmental gradient where the rains start at different times each year south-to-north of Nigeria. To test this, we modelled moult progress just before the rains across 15 localities 6°–13° N as a function of the onset of the wet season among localities. As predicted, moult progressed further in localities with earlier wet seasons, confirming that the onset of moult is timed to the onset of the wet season in each locality despite weak breeding seasonality in the Common Bulbul. This strategy may evolve to maintain optimal annual cycle routine in seasonal environments where breeding is prone to unpredictable local perturbations like nest predation. It may, however, be less obvious in temperate systems where all annual cycle stages are seasonally constrained, but it may help with explaining the high frequency of breeding–moult overlaps in tropical birds.
Highlights
Life history traits should be timed to coincide with suitable environmental conditions or at different times from conflicting traits that may compete for common resources (Barta et al 2006; McNamara and Houston 2008; Wingfield 2008; Visser et al 2010; McNamara et al 2011)
The seasonality of many such traits and the annual routine of many organisms are well documented, identifying the environmental cues that govern seasonality is difficult because the seasonality of environmental conditions and that of annual cycle stages are correlated in time (Dawson 2008)
In central Nigeria, for example, we observed that Common Bulbuls Pycnonotus barbatus moult in the wet season at the population level despite showing weak breeding seasonality, and we concluded that moult was more sensitive to environmental seasonality than breeding (Nwaogu et al 2019)
Summary
Life history traits should be timed to coincide with suitable environmental conditions or at different times from conflicting traits that may compete for common resources (Barta et al 2006; McNamara and Houston 2008; Wingfield 2008; Visser et al 2010; McNamara et al 2011). In birds, breeding and moult are often timed to spring and summer conditions in temperate environments and to the wet season in most tropical environments (Baker 1939; Hau 2001; Repenning and Fontana 2011; Mares et al 2017). It is, unclear whether the timing of each annual cycle event is related to independent cues, flexible to the timing of other events or is dominated by the need to time the most important annual life history event to a seasonal optimum. We predicted that moult progress will correlate with the timing of the onset of the wet season among localities
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