Up until relatively recently, it was hard to study the same population of migratory waders in both its breeding area and its wintering grounds. Ringed birds established links between different countries but to follow a group of individuals through a complete annual cycle was nearly impossible. Geolocators, and more recently satellite tracking, are starting to enable scientists to piece together whole stories.
Camilo Carneiro and colleagues from the University of Aveiro in Portugal (Dep. Biology & CESAM) and the University of Iceland (South Iceland Research Centre) have been tracking Whimbrel travelling between Iceland and Africa for nearly ten years, using geolocators. In the latest paper to come out of this research they investigated carry-over effects; do conditions experienced in wintering locations affect breeding success?
How might carry-over effects work?
The conditions experienced during one stage of a migrant’s annual cycle may affect their performance in subsequent stages. Perhaps the resources available at a wintering site might affect the timing of spring departure and whether an individual has to stop off to refuel? In turn, such individual differences may be apparent in individuals’ arrival dates in the breeding area, and the condition they are in might affect laying date, clutch size, egg weight, etc?
Icelandic Whimbrel spend the wintering season anywhere between south-west Europe and the tropical coastal areas of West African countries such as Benin and Togo. Despite this huge non-breeding range, individuals are highly philopatric, travelling between the same breeding area and the same restricted wintering site on an annual basis, perhaps for twenty or more years. A Whimbrel flying to the Bijagós Archipelago of Guinea-Bissau covers nearly 6000 km, in the autumn, whereas a bird that only travels to the Tejo (Tagus) Estuary of Portugal flies not much more than half as far (see map). The ‘winter’ conditions they experience are completely different; short temperate days in Portugal or tropical heat in the mangroves in Guinea Bissau.
Camilo and colleagues were able to study Whimbrel in different wintering locations, in order to understand the conditions that are experienced by breeding birds from these areas. They measured annual return rates for birds that had flown different distances and experienced different conditions in the non-breeding season. Do Tejo birds, spending the non-breeding season in the coldest part of the wintering range, have a lower apparent chance of survival? Do those that make it through a Portuguese winter return to Iceland earlier and thereby increase their chance of breeding successfully?
Life on the wintering grounds
Camilo Carneiro has studied wintering Whimbrel in three sites – the Tejo Estuary (Portugal), the Banc d’Arguin (Mauritania) and the Bijagós Archipelago (Guinea-Bissau). Birds in the three sites experience very different conditions between the start of September and the end of March, as discussed in the paper and illustrated in the table below.
Hundreds of observations of individual Whimbrel and flocks provided information on feeding rates, diet and foraging time. Comparable food items were collected from the mud/sand substrates and the energetic values were calculated in the laboratory. Together, these data enabled a calculation of energetic intake. The Net Energetic Intake Rate varied markedly. The figure for the Bijagós is 3.9 times that of the Tejo and 1.4 times that of the Banc d’Arguin.
Birds have a basic running cost – the Basal Metabolic Rate – which is related to the size of the individual and ambient conditions it experiences. Those wintering in areas where they experience periods of colder and windier weather lose more heat and hence need more energy. The BMR was calculated as 2.17, 2.29 and 2.51 Watts for individuals wintering in the Bijagós, Banc d’Arguin and Tejo, respectively, showing that there are higher ‘running costs’ in northern sites. Whimbrels in the Bijagós never incurred energetic costs above BMR, whereas those in the Banc d’Arguin and Tejo had additional energetic costs on 20.6% and 9.7% of the winter days, respectively.
The daily energetic balance differed hugely. Whimbrels in the Bijagós experiencing an average energetic surplus of about 700 kJ/day, followed by 420 kJ/day in Banc d’Arguin and just 11 kJ/day in the Tejo. It should be noted that these figures are based only on day-time feeding.
Returning to Iceland
The research team has shown that Whimbrel can either fly directly to Iceland or stop off and refuel. It is thought that between 80% and 90% of journeys include a stop-over, typically in Ireland or western Britain. Direct flight takes four or five days. (see summary of previous papers and blogs below).
Whimbrels arrive back in Iceland between the end of April and late May, quickly taking up territories unless there is snow cover. Regular visits to the main study area in the Southern Lowlands helped to ascertain which colour-ringed birds had returned when. Nests are found, and eggs are measured and ‘floated’, to estimate the laying date.
During incubation, attempts were made to catch marked individuals that were carrying leg-mounted geolocators. Adults trapped on the nest are measured, unringed birds are marked and three to five feathers are removed from the breast. These feathers will have been grown in the bird’s wintering area and carry an isotopic signature from that region.
Using stable isotope analyses of the breast feathers, ground-truthed by birds tracked using geolocators, Camilo managed to assign the winter location to 180 Whimbrels. 159 had flown from the tropical region (which includes Bijagós), while 20 had spent the winter in the arid region (which includes Banc d’Arguin) and one in the temperate region (Tejo). When linking the wintering region to breeding phenology and investment, the research team found that:
- There were no differences in the size of the birds returning from the tropical and arid regions.
- There was no difference between the probabilities of a bird successfully returning from the tropical and arid regions.
- The timing of nesting and the volume of the eggs that were laid by females was not different for birds from the tropical and arid regions.
Where to spend winter?
Only one marked bird definitely wintered in temperate southwest Europe, which is not surprising given that there are not large flocks of Whimbrel in the estuaries of this area. This bird was excluded from the analyses but we know that it will have travelled much less far than birds wintering in Africa and experienced winter conditions in which it could barely meet its daily energy requirements.
Individuals wintering in the arid region, including birds in the Banc d’Arguin, travelled a lot further than birds wintering in southwest Europe. These birds had an expected surplus of 420 kJ per day on an average day but strong winds meant that there were 20% of winter days in which conditions were sub-optimal.
Individuals wintering in the Tropical group, including birds in the Bijagós, travelled 900 km further than the Arid group but found more predictable weather conditions, achieving an estimated spare energy capacity of 700 kJ per day, without days with energetic costs above BMR. The authors point out that this energy surplus will likely be needed during long periods of moult and to fuel spring migration.
The authors conclude that any costs associated with having to fly further to reach the tropical region are compensated for by benign conditions. This does not mean that an individual bird makes a choice between Tejo, Banc d’Arguin and Bijagós. Happenstance may determine where a juvenile ends up in its first winter and philopatry means that, if alive, it continues to spend subsequent winters in the same area. Presumably the risks incurred by flying further (to Bijagós) balance out the risks incurred by wintering in a less predictable environment.
Very few Whimbrel spend the winter on the Tejo, in Portugal, and calculations in the paper suggest that there is a high risk of not being able to find enough food. This would probably translate into high mortality and explain low numbers.
Although no carry-over effects were found, the authors discuss ways in which they may show up in other traits. There is an interesting discussion as to how carry-over effects might link experiences in the wintering grounds to breeding output in the next breeding season. Amongst other things, the authors suggest that differences among individuals using different wintering sites may only become evident if assessed over several years. We know that conditions are less benign in Banc d’Arguin, for instance. Perhaps there are years when conditions are bad enough for long enough to influence survival or body condition in spring. In such a year, there could be impacts on the ability to migrate, an increased likelihood of dying during migration or delayed breeding. It is possible that longer-term studies will pick up differences in return rates and/or breeding success for birds wintering in different areas? We shall see.
Linking range-wide energetic trade-offs to breeding performance in a long-distance migrant Camilo Carneiro, Tómas G. Gunnarsson, Verónica Méndez, Amadeu M.V.M. Soares & José A. Alves
Previous research on Icelandic Whimbrel
Whimbrels on the move summarised the movements of Icelandic birds, based on reports of ringed and colour-ringed individuals. In the paper upon which the blog was based (Gunnarsson & Guðmundsson) there was a strong suggestion that birds only stop off in Britain & Ireland on the way north. Geolocator-based research by Alves et al showed that at least some birds were flying straight from Iceland to West Africa and that these sea-crossings could be very rapid.
Migrations to and from Africa were investigated further in a paper by Camilo Carneiro et al that was summarised in Iceland to Africa, non-stop. More recently, papers by the same team have shown that the most consistent point of the annual migration story is departure from Africa and discussed the links between weather and phenology. These two papers have appeared as the WaderTales blogs – Whimbrel: time to leave and A Rhapsody of Whimbrel.
The following WaderTales blogs all consider how migratory behaviour might affect breeding season success, although without the direct measurements for individuals that have been carried out in the Whimbrel study.
Overtaking on migration shows that potential costs of migrating further can be overcome by undertaking early spring migration to staging sites that are closer to breeding areas.
Travel advice for Sanderling summarises research to understand the pros and cons of spending the non-breeding season in widely different locations.
Gap years for sandpipers is based upon a Peruvian Semipalmated Sandpiper paper that investigates the survival advantage of not migrating north to breed in a particular year.
WaderTales blogs are written by Graham Appleton (@GrahamFAppleton) to celebrate waders and wader research. Many of the articles are based on published papers, with the aim of making shorebird science available to a broader audience.