There’s a lot to fit into twelve months if you’re a Whimbrel. In the last paper from his PhD, Camilo Carneiro assesses whether Icelandic Whimbrel can always manage to complete the annual cycle of migrate-breed-fatten-migrate-moult-fatten in just 365 days.
What happens if a pair of Whimbrel loses a first clutch and a successful, late breeding attempt delays departure from Iceland, for instance? Is there the flexibility to make up for lost time during a west-African winter? Some answers are provided in a paper by Camilo Carneiro et al in The American Naturalist.
Facing the consequences
We know that large waders sometimes take a year off from breeding, as discussed in Teenage waders, suggesting that they may not always have the resources they need to breed every year. This blog, based on a paper by the Bird Ecology Lab team in Chile, tells the tale of a few Hudsonian Godwits that headed for the pampa grasslands of Argentina in spring, instead of migrating to Alaska.
As discussed in Gap years for sandpipers, taking a year off may make no difference to expected lifetime breeding output and could be more common when individuals spend the non-breeding season a long way from nesting areas or in poor quality sites. If Iceland’s Whimbrel are able to compensate for any delays they face during the course of the annual cycle then perhaps that suggests that all is well for this population that migrates all the way to West Africa at the end of the summer breeding season?
The story so far
Whimbrel only spend three months of the year in Iceland, with the rest of the time spent some 6000 km further south. In July or August, an adult will exchange the sparsely vegetated river plains and night-time frosts of Iceland for mud, mangroves and tropical temperatures. The diet changes too, from terrestrial invertebrates and crowberry to crabs.
Camilo Carneiro started a PhD on Iceland’s breeding Whimbrel in 2015, continuing the work of his supervisors, Tómas Gunnarsson of the University of Iceland and José Alves from the University of Aveiro (Portugal). Using geolocator tags he investigated the capacity for Whimbrels to undertake non-stop journeys, demonstrating that autumn migration was generally direct but spring migration for most birds included a stop, often in Ireland or the UK. You can read more in Iceland to Africa, non-stop. Birds that need to (or choose to) take a break delay their arrival in Iceland by about ten days.
Subsequent papers by the same team have shown that the most consistent point of the annual migration story is departure from Africa and shown links between weather and phenology. These two papers have been covered in the WaderTales blogs Whimbrel: time to leave and A Rhapsody of Whimbrel. In an attempt to discover the best places to spend the winter months, Camilo analysed tag and colour-ring data to work out links between conditions experienced in wintering locations and subsequent breeding success, as discussed in the blog Winter conditions for Whimbrel.
Seven years of data
To investigate how migratory animals navigate their annual schedule, and where and when they can make adjustments to their timings, Camilo Carneiro and his colleagues used annual-cycle data of 38 Icelandic whimbrels tracked over 7 years. They asked three questions:
- Does the change in the timing of one event in the annual calendar, such as a late breeding season, affect the timing of subsequent events, perhaps with further down-stream domino effects?
- Can individuals compensate for delays, on migration for instance, by spending less time on the next stage of the annual cycle, e.g. by reducing a stop-over?
- Are there potential fitness consequences? Do birds that are subject to delays breed later? In waders, earlier chicks are more likely to recruit to the breeding population so being just a few days late returning to Iceland may have consequences.
During the period 2012 to 2018, a total of 78 geolocators were deployed on Whimbrel breeding in Southern Iceland. The device was attached to a leg-flag in one year (see picture) and usually collected in the subsequent breeding season. In most cases, a replacement geolocator was fitted, in order to collect further data on that individual. Unsurprisingly, it became harder to catch tagged birds over time, as birds learned to recognise nest traps and research vehicles. The fact that so much valuable information was collected from the same birds is testament to Camilo’s patience. Sixty-six geolocators were retrieved from 39 individuals. Birds could only be caught when incubating a full clutch of eggs so nest losses due to predation affected the likelihood of recapture.
Please see the paper in The American Naturalist for full details of the methods used to collect data on breeding success and for interpretation of data collected using geolocators.
Before looking at the results in the paper, here’s an example of data collected using geolocators, for two years in the life of YY-LL (Yellow Yellow – Lime Lime), pictured alongside.
In 2015, YY-LL nested successfully and left Iceland on 16th August. After four days of direct flight, he reached Guinea-Conakry. His nesting attempt in 2017 was unsuccessful and he migrated south a little earlier, on 5th August, again taking four days to fly 6000 km.
In the springs of both 2016 and 2018, YY-LL left the winter grounds on 22nd April, arriving in Ireland on 26th April in 2016 and on 25th April in 2018. Migration from Ireland to Iceland took place between 5th and 7th May in 2016 and between 7th May and 9th May in 2018.
YY-LL hints at strong consistency of spring migration timing, independent of nest success.
The 38 tagged Whimbrel provided information about 76 autumn migrations and 60 spring migrations. Most of the birds (89%) spent the winter between Senegal and Sierra Leone, particularly in Guinea-Bissau and Guinea, with one bird in Portugal and the rest in coastal northwest Africa.
As is usual in waders, females left Iceland before males, the difference being typically around six days. Although failed breeders did depart earlier than birds that successfully reared chicks, the difference was not great, again averaging around six days.
There was no suggestion that birds that left Iceland late in the season ended up on a delayed schedule for return to Iceland in the subsequent spring. This implies that resources in Africa were sufficient to ‘catch up’ with earlier birds, despite the need to complete a full moult and to prepare for another 6000 km migration.
As indicated in previous papers (and in the blogs mentioned above) spring departure dates of tagged birds from Africa were not different for different countries or for different sexes. However, birds that stopped in Europe tended to leave Africa earlier (19th April on average) than those that made direct spring flights (30th April). These latter birds tended to arrive in Iceland about a week earlier than birds on a two-stage migration, representing a neat, overtake manoeuvre! Typically, males arrived in Iceland about a week earlier than females.
For the sample of tagged birds, neither the autumn departure date from Iceland nor wintering location had any apparent effect on the arrival date in the next spring.
There seems to be a strong signal that, just as YY-LL did, an individual Whimbrel can make up for any delays incurred, with birds that arrive at a location later spending less time there, whether that be a wintering site or a spring stopover location. Females tended to spend longer at spring stopover locations, which ties in with the earlier arrival of males in Iceland. The graph alongside illustrates these two points. Birds that left Africa earliest spent more than 15 days at stopover sites but birds on a later schedule stopped off for as few as 6 days. Triangles represent males and squares are females.
Over the course of a year
Putting this all together, Camilo and his colleagues found that individuals appear to use the wintering sites to compensate for delays, these mostly having been associated with a successful previous breeding season. The wintering season is up to 38 weeks long so a Whimbrel that heads south a little late has plenty of time in which to catch up with earlier birds. The timings for other wader populations that spend shorter periods in wintering locations may be more constrained, given that post-breeding moult might take 20 weeks and the time to fatten up for migration can add an extra seven weeks.
Once a bird leaves Africa, it is harder to compensate for delays, although attempts are made to do so, with later individuals stopping for shorter times at spring stop-over sites and then nesting shortly after arrival in Iceland. Data in the paper suggest that it is not possible to catch up completely, before the start of the breeding season, if time is lost on the way north. Given the known link between lay-date and nesting success, these spring delays may have consequences for productivity and reduce the capacity to re-nest following clutch loss.
Camilo’s research suggests that adult Whimbrel in the south of Iceland have the capacity to make up for any delays that they face during the annual cycle. The same may not be true for other large waders, the populations of which are mostly in decline. The blog Why are we losing out large waders? reminds us that two curlew species are thought to be extinct or on the verge of extinction and that most of the rest are in trouble.
All is not necessarily well for Iceland’s waders either. Two 2022 papers by Aldís Erna Pálsdottir, looking at the effects of forestry and power-lines, suggest that the Whimbrel is one of the wader species most seriously impacted by ongoing changes to the Icelandic landscape, as discussed in Power-lines and breeding waders and Iceland’s waders need a strategic forestry plan. These pressures seem to be reflected in poorer breeding output, as suggested by counts of family parties of Whimbrel made by Tómas Gunnarsson and colleagues in annual June and July surveys.
Camilo’s paper may indicate that adult Whimbrel can cope with all that life throws at them but if they cannot raise enough chicks the species will still be in trouble. With financial support from the Icelandic Centre for Research, Camilo is now studying the challenges that chicks face, as they prepare for their first migration from Iceland to Africa.
Here’s the link to the paper:
Annual schedule adjustment by a long-distance migratory bird. Camilo Carneiro, Tómas G. Gunnarsson & José A. Alves. The American Naturalist. doi.org/10.1086/722566
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.