Jennifer A. Gill, José A. Alves, William J. Sutherland, Graham F. Appleton, Peter M. Potts and Tómas G. Gunnarsson. 2013. “Why is timing of bird migration advancing when individuals are not?” Proc. B. Vol. 281, no. 1774.
Phenological responses have been used as one of the major indicators of climate change. The timing of flowering and fruiting, the return of migrant birds and insects from winter habitats are easily and often measured, and records going back decades or centuries sometimes exist. Most importantly, shifts in phenological indicators are some of the strongest connections between rising temperatures and biological and ecological responses (
for example). There is plenty of
evidence, for example, that some migrant bird species are returning to their breeding grounds earlier than ever. These migratory birds may be responding (via migration timing) to warming temperatures in several ways: there may be plasticity or flexibility in individual timing of migration which allows them to respond to changing temperature cues; or species may also show adaptation via changes in the frequency of individuals with different migratory timings (microevolution). In cases where migratory species are responding to climate change, distinguishing the mechanisms allowing them to do so is surprisingly hard. Early arrival of migratory bird species is often explained as being due to individual plasticity or flexibility in “choosing” the date of migration, but the majority of studies of this phenomenon include little or no information about individual behaviour, only changes in the mean date of arrival for the entire population.
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For this reason, Gill
et al. looked at individual, rather than average population, arrival dates for Icelandic black-tailed godwits in south Iceland. Icelandic black-tailed godwits (“godwits” for the sake of brevity) have shown significant advances in the last 20 years in the timing of their spring arrival to the shores of Iceland, and these advances appear to relate to increasing temperatures.
The population has also been banded such that 1-2% can be individually identified and tracked throughout their migratory range. Although only adults (of unknown age) were banded at the start of the experiment in 1999, recently chicks have also been banded and released and so a wide range of demographic classes are included with the banded birds.
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From Gill et al. 2013. |
When Gill
et al. looked at date of arrival across 14 years for each individual, their results were surprisingly clear and cohesive. As previously reported, the population mean date of arrival in South Iceland had advanced as much as 2 weeks. But, this advance is not reflected in individual timing of arrivals over that same period – if a bird tends to arrive on a given day, they will continue to arrive on approximately that day every year, independent of temperature conditions. Instead, the population trend appears to be driven entirely by birds born in recent years – young individuals (recently hatched) tend to have arrival dates much earlier than older individuals. At least for the godwits, population wide trends in migration dates are actually driven by only a subset of the individuals.
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From Gill et al. (2013) |
Often it is assumed that migratory birds are responding to warming temperatures on an individual level: individuals respond to changing cues, resulting in shifts in arrival date. This study suggests otherwise, and finds that the important mechanism is not individual plasticity or microevolution but rather related to demographic shifts in arrival time. As to why younger birds arrive earlier, it is not clear, but may relate to the observation that nest building and hatching dates are also advancing. It may be that natal conditions are important – the authors observed a variety of possibly inter-related changes such that hatching dates are advancing and chick sizes are increasing, and the suggestion that mortality rates of later arriving individuals may also be higher. "Environmentally induced advances in arrival dates of recruits could operate through: (i) carry-over effects of changing natal conditions, (ii) changing patterns of mortality of individuals with differing arrival times, or (iii) arrival times being initially determined by conditions in the year of recruitment and individuals repeating those timings thereafter."
These results make some predictions about which populations of migratory birds might have the most ability to respond to warming climate - most likely those with shorter migratory distances, shorter times to reproduction and shorter-lifespans (hence decreasing the lag-time required for the population to catch up to temperature). It may also have relevance for other non-bird species that also rely on careful timing between phenology and temperature. Correspondingly, it suggests limitations - if individual behaviour is so inflexible and constrained, our hopes that some species may respond to climate change with behavioural changes seem far to simplistic.
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