Whimbrels arrive in Iceland

There is something magical about standing in Eyrarbakki in South Iceland, in spring, watching small flocks of Whimbrel come in over the sea. Thanks to geolocators and satellite tags (about which more later) we know that these amazing waders will have been in the air for around five days, since leaving the west coast of Africa. Looking down at the seaweed-covered rocks on the morning of 27th April 2022, we could pick out small groups of new arrivals. Most were resting but one bird was looking for crabs, just as it had been doing in West Africa just a few days previously. A few birds headed off inland while we were watching, making the distinctive seven-note whistle as they left. It sounded almost like a cry of “made it”!

Incoming Whimbrel: Tómas Gunnarsson

Setting the scene

We had been here before. On 22nd April 2008, I did a live broadcast for the BBC Radio 4 programme World on the Move from this very spot, when I described visible migration to Brett Westwood. On that day, we could see Purple Sandpipers and Turnstone feeding on the tide edge, White Wagtails and Meadow Pipits, newly arrived from Iberia, and small flocks of Golden Plover flying in low over the sea. During the programme, a gaggle of 35 Pink-footed Geese flew in strongly from Britain, as did four Arctic Skuas that had spent winter in the Southern Hemisphere.

On 27th April 2022, the skies were not as busy as they had been fourteen years previously. High pressure over the British Isles and northerly winds across the Atlantic seemed largely to have pressed ‘pause’ on migration from Britain & Ireland. There were newly-arrived White Wagtails, Meadow Pipits and Black-headed Gulls feeding on insects emerging from the banks of rotting seaweed, but the stars were definitely the Whimbrel.

These banks of seaweed will be even more important in May, when there will be hundreds of waders refuelling for the next leg of their journeys, with Sanderling and Dunlin on their way to Greenland and some Knot and Turnstone perhaps flying as far as northeast Canada. When the tide is high enough to wash maggots out of the seaweed, you can sometimes see Red-necked Phalaropes along the tide-line, spinning around and picking food off the surface of the water.

Whimbrel migration

We knew that the Whimbrel were on their way because Global Flyway Network had published a map showing the location of ‘Acuno’, a bird that is wearing a satellite tag that was put on in the Bijagós archipelago of Guinea Bissau. It had been logged west and north of Ireland on the previous evening, already 6000 km and 4 days into its migration. Doing the sums, it seems unlikely that it could have reached Eyrarbakki by the time that we were there but perhaps it flew past soon after, sending signals back to mission control that would confirm arrival**.

** An hour after I published this blog, Acuno was found to have diverted to the Faeroes. It may breed there – as many Whimbrel do – or if could have run out of fuel and landed there, to put on some extra grammes of fat. We’ll see whether it resumes migration.

** Three weeks later, Acuno flew to Iceland. How much will it have been disadvantaged by giving up on direct flight?

Only a minority of Whimbrel fly straight from Africa to Iceland in spring although almost all fly directly south in the late summer. Most individuals spend late April and early May in Ireland, the UK or on the west coast of mainland Europe. Ireland is by far the most important staging area. The individuals on Eyrarbakki beach may have been tired but there could be advantages to being an early bird. See Time to nest again? based on a paper by Morrison et al.

There are several WaderTales blogs about Icelandic Whimbrels:

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 latest blog about this research is Winter conditions for Whimbrel, based on a paper that assesses the influence of winter conditions on subsequent breeding performance.

Searching for Black-tailed Godwits

We had seen a flock of eight Whimbrel, earlier in the morning, when we were checking fields for colour-ringed Black-tailed Godwits. The Whimbrel were gliding into land, about five km from the coast, before seeming to melt into a patch of rough grassland, bleached after the winter snows.

We did not pay the Whimbrel much attention as, on the other side of the road, there were Black-tailed Godwits probing for worms in silage fields that were already green, after a few days of warmth and the liberal addition of fertilizer. This is our target species during spring trips to Iceland. We have discovered that the arrival time of individuals is remarkably consistent from year to year, which initially seemed surprising, given that migration appears to be getting earlier. There is more about this in Why is spring migration getting earlier? based on a paper in Proceedings of the Royal Society B.

One of the fascinating things about visiting Iceland is that no two years are the same. 2022 has been a dry, warm spring, with northerly winds potentially delaying migration from Britain and Ireland, as mentioned earlier. Early-arriving Black-tailed Godwits that were wearing colour-rings were birds that winter in Portugal and France and migrate via the Netherlands. There was a period of helpful winds for these early birds that fly further but get to Iceland earlier. This strategy is discussed in the blog Overtaking on Migration, based on a paper in Oikos.

Looking forwards

The short Icelandic summer provides fantastic conditions for these waders to raise their chicks, although there are concerns as to how agricultural development, increased forestry and infrastructures will affect these species in the future. In June, in just two months’ time, adults will cross the Atlantic. Black-tailed Godwits head for the British Isles and the west coast of continental Europe and Whimbrel will return to West Africa. By August, the next generation will be preparing for the journey south and we will be here in future springs to monitor their return to Iceland.


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.

Navigating a vast ocean

The journeys that shorebirds make, as they cross the oceans of the globe, are truly remarkable. Individual birds demonstrate amazing endurance and navigational expertise while in the air for days at a time. Satellite transmitters are providing opportunities to understand how shorebirds refine their flight plans in responses to the wind patterns they encounter. In a paper in Movement Ecology by Jenny Linscott and colleagues, we join the northward flight of Hudsonian Godwits across 7,000 km of the Pacific Ocean and the Gulf of Mexico.

Coping with wind

Conditions encountered en route can dramatically impact the energy that migratory species spend on movement. To complete their journeys across barriers such as oceans, migratory birds need to manage energetic costs by adapting to the wind conditions they encounter. It’s a dynamic environment; no two years are the same and there is often little relationship between local conditions upon departure and what lies ahead, as for instance discussed in the WaderTales blog Rhapsody of Whimbrel.

Migrating Hudsonian Godwits are heading for Alaska

If an individual bird has full knowledge of its location at all times and a fixed destination point in mind, then perhaps it can adjust its airspeed and direction of flight so that it can completely compensate for any lateral drift that is caused by the wind systems en route. Alternatively, it might accept the drift that it experiences or only partly compensate, perhaps gambling that there may be systems ahead that might cause drift in the other direction. It may even over-compensate for the wind patterns it experiences at a particular time, which may be a good idea if there is more of the same weather to come. The authors of the Hudsonian Godwit study explain how these different approaches may integrate to create a movement strategy and summarise what has been found in other studies. Interestingly, it has been suggested that complete compensation (continually adjusting the flight-plan) might not be possible for trans-oceanic flights because there are no landmarks to use as reference points.

Hudsonian Godwits spend the austral summer in coastal Chile and Argentina and migrate northwards in spring, through the midcontinental United States, to breed in subarctic Alaska and Canada. For birds taking off from Chiloé Island in Chile, the first part of the journey takes them across the open Pacific Ocean, the relatively narrow landmass of Central America and the Gulf of Mexico. Godwits making this flight have few or no opportunities to stop, and they traverse several global wind regimes that differ in directionality and strength along the way. The authors predicted that birds would experience drift during their journeys, especially over the featureless open ocean, and increase compensation as they approach North America. Given the vast distances and time spent in flight, it might be expected that the winds experienced during flight should influence the points where individuals cross the coast into North American airspace.

Flock of Hudsonian Godwits on Chiloé Island in Chile – a long way from Alaska

Tracking transoceanic journeys

During the springs of 2019, 2020 and 2021, Jenny Linscott used satellite tracking devices to follow Hudsonian Godwits, as they migrated northward across more than 7,000 km between Chiloé Island, Chile and the northern coast of the Gulf of Mexico.

Prior to migratory departure, two types of solar-powered satellite transmitters were deployed on a total of 54 adults. Technical details are provided in the paper, alongside information as to how the data were filtered and interpreted. This paper focuses on the journey across the Pacific Ocean and the Gulf of Mexico, as birds head towards a relatively narrow refuelling area in North America (the eastern parts of Kansas, Nebraska, South Dakota, and North Dakota).

For any two points along each bird’s route, between Chiloe and the coast of the Gulf of Mexico, it was possible to calculate distance travelled, ground speed, turning angle, and heading. Having removed low-quality data, the team were left with 29 tracks and 689 locations with which to work. They linked each location with the range of possible wind conditions that godwits were likely to be experiencing at the time, depending upon flight height. They then “calculated the total magnitude and mathematical direction of the wind flow for each location at each altitude using vector trigonometry”. Previous studies have shown that migrating waders change altitude in order to find better wind conditions, allowing the team to assign the most likely wind conditions to each godwit location. Please see the paper for details of the modelling and assumption-testing processes.

Results

A total of 24 complete and 5 partial northward journeys were collected from birds migrating north, including repeat tracks from two individuals, which were followed for three years. The sample size was reduced by device failure/malfunctions, presumably some mortality, and by eight birds that oversummered in Argentina (see Teenage Waders and Gap year for sandpipers for information about oversummering shorebirds).  The 25 birds in the study comprised 14 males and 11 females.

  • Godwits for which there were complete tracks undertook continuous flights lasting about six days, covering an average of 8,361 km, before making their first stops.
  • Godwit ground speeds were best predicted by a strategy in which individuals flew at the altitude offering the most wind support in the preferred direction of movement, but were restricted to altitudes at or below 3000 m. It seems likely that the godwits were mostly flying between 100 m and 750 m above sea level.
  • Tracked godwits travelled along paths which showed a close match to a Great Circle line from Chiloé to the North American target area.
  • When flight behaviours were analysed, full compensation was the most frequent behaviour, accounting for 41.1% of all observed flight segments. Fewer segments were associated with partial compensation (23.5%), tail winds (8.1%), full drifting (9.4%), or overcompensation (16.0%).
  • The prevalence of full compensation remained constant across wind conditions. For example, full compensation was the dominant behaviour under crosswinds to the east (32.0%) and west (45.3%). Full compensation was also prevalent across regions, comprising the largest proportion of behaviours exhibited over the Pacific Ocean (45.1%), while crossing Central America (23.9%), and in the Gulf of Mexico (31.4%).
  • There was considerable variation in migration patterns over the Gulf of Mexico and it is suggested that some birds may have been running out of fuel and heading for the nearest land.
  • One individual tracked repeatedly over three years completed its crossing of the Gulf of Mexico in central Texas every year. The other bird tracked for three years had no specific point at which it crossed the coast but always ended up in the same spring staging area.

Jenny Linscott and colleagues found little support for their prediction that godwits would tolerate more drift early in their flight and gradually begin to increase compensation as they approached North America. Instead, both fully supported flight (benefiting from tail winds) and full compensation were common soon after leaving Chiloé. Compensation did not increase with distance travelled, was not constrained during flight over open ocean, and did not influence where an individual ultimately crossed over the northern coast of the Gulf of Mexico, at the end of this flight. Instead, the team found a strong preference for full compensation throughout godwit flight paths. Birds ‘knew’ where they were along their route and could judge how to adjust their headings so as to compensate for the drift they were experiencing at any given time. The paper’s Discussion includes more detailed consideration as to how compensation appeared to operate in different parts of the journey and in different wind conditions.

How do shorebirds, flying over the vastness of an ocean ‘know’ where they are, with no island landmarks? Are individuals within migrating flocks picking up on changes in temperature or humidity, that mark passage through broad wind regimes, can they navigate with reference to sun position and stars, are there magnetic cues, or can they interpret surface swell patterns? Perhaps it’s a mixture of several of these skills? It’s pretty amazing!

Learning more

Jenny Linscott and her colleagues found that fully compensating for wind displacement appears to be a critical strategy for Hudsonian Godwits making a long-distance, transoceanic flight. While godwits often followed wind flow in the early stages of this journey, they nonetheless engaged in full compensation more frequently than any other behaviour during the entirety of the flight, across a vast and apparently featureless ocean. These continuous adjustments help to make sure that birds can fly extremely long distances without running out of energy. The team wonder how well future generations will cope with changing wind systems over warmer seas.

The full paper can be found here:

Compensation for wind drift prevails for a shorebird on a long-distance, transoceanic flight. Jennifer A. Linscott, Juan G. Navedo, Sarah J. Clements, Jason P. Loghry, Jorge Ruiz, Bart M. Ballard, Mitch D. Weegman, and Nathan R. Senner. Movement Ecologyhttps://doi.org/10.1186/s40462-022-00310-z

The Hudsonian Godwits studied here are flying 7,000 km north across the Pacific, but this pales into insignificance when compared to Bar-tailed Godwits that commute between Alaska and New Zealand. The Pacific was once considered a barrier to migration but it is increasingly seen as a conveyor belt. There is an excellent review article by Theunis Piersma and colleagues in Ornithology, the title of which explains its content:

The Pacific as the world’s greatest theater of bird migration: Extreme flights spark questions about physiological capabilities, behavior, and the evolution of migratory pathways. Theunis Piersma, Robert E Gill, Jr, Daniel R Ruthrauff, Christopher G Guglielmo, Jesse R Conklin and Colleen M Handel. Ornithology,  https://doi.org/10.1093/ornithology/ukab086


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.

How many shorebirds use the East Asian-Australasian Flyway?

The level of threat facing a shorebird species is assessed using a set of criteria that tries to answer three key questions: How many breeding pairs are there? How quickly are numbers falling? and How restricted is the distribution? In a paper published in Ibis at the end of 2021, Birgita Hansen and colleagues produced population estimates for 37 species of shorebird on the East Asian-Australasian Flyway, for which the non-breeding distribution includes Australia and New Zealand. This new study shows that there are at least nine million shorebirds on this flyway. How many more would there have been ten, twenty or fifty years ago?

Population estimation

More comprehensive data are now available for countries such as Myanmar

This is a good time to attempt to assess population sizes of shorebird species on the East Asian-Australasian Flyway because the quality of the data available is so much better than it was even ten years ago. Increased cooperation across borders, fostered through individual efforts by concerned birdwatchers, by conservation NGOs and through bodies such as the East Asian-Australasian Flyway Partnership, has delivered reliable counts of shorebirds in previously inaccessible areas, especially around the Yellow Sea.

The East Asian-Australasian Flyway covers an area of 85 million square kilometres, from Russia and Alaska, in the north, through to Australia and New Zealand in the south. Birgita Hansen and colleagues have revised the population estimates for 37 migratory shorebird species protected under Australian national environmental legislation. Population estimates were generated by:

  • Summarising existing count data in the non-breeding range
  • Extrapolating the data to try to include the numbers of birds using uncounted areas and
  • Modelling abundance on the basis of estimates of breeding range and density.

This was not just a number-crunching exercise; expert opinion was sought, especially to refine estimates for species with low data quality.

Flocks of roosting shorebirds on the white sand of Eighty Mile Beach in Western Australia

The main count data available to the authors came from the Australian National Shorebird Monitoring program (formerly Shorebirds 2020), the Queensland Wader Study Group, the Ornithological Society of New Zealand, and the Asian Waterbird Census. Two major strengths of the paper are a clear explanation of the methods used to create the population estimates and open consideration of the quality of the estimates. The authors explain how it is possible to produce population estimates in areas where there are significant gaps in the data. These features will be important to teams making future assessments of numbers.

The Hansen paper builds on previous estimates by Bamford et al, in a paper published in 2008, that used data collected between 1986 and 2000, and through an expert-assessment process (Wetlands International 2012). However, the various estimates have been derived in different ways and cannot directly be compared.

There is a summary of the Hansen population estimates on the Department of the Environment (Australia) website. Context and caveats are provided in the new paper.

How many shorebirds?

How many shorebirds (and terns)?

Species that form coastal flocks in a relatively well-established set of sites are easier to census than others that are spread thinly or across a range of habitats. Flyway counts of Curlew Sandpiper are thought to include almost all of the sites used by the population, so a total of 85,086 is adjusted to an estimate of 90,000 to account for gaps. For Latham’s Snipe, the flyway estimate is 35,000, based on the breeding range and density estimates, but actual flyway counts only total 1124. For Curlew Sandpipers, non-breeding season numbers appear to be reliable indicators of population size but for Latham’s Snipe breeding season estimates are more realistic. Variability in data quality is not unusual; the British & Irish estimates of winter numbers of Common Snipe, Jack Snipe, Eurasian Woodcock, Golden Plover and Northern Lapwing are similarly imprecise, as discussed in Do population estimates matter? and Ireland’s wintering waders.

Estimates of population size obtained through analyses of breeding ranges and density indicated that non-breeding counts for 18 species in the Hansen study were relatively uncertain. Breeding ground estimates were considered the best available data for ten species that mostly use poorly-surveyed freshwater and marine habitats, especially in south-east Asia, or are thinly distributed on the open coastline in the non-breeding season. The WaderTales blog Waders on the coast reflects on the latter issue for UK waders.

Birds at risk

Prioritisation of conservation action for shorebirds is underpinned by the threat levels assigned to them. This means that it is probably easier to access funding for Critically Endangered Spoon-billed Sandpipers than for Near Threatened Curlew Sandpipers. There are three categories indicating imminent risk of extinction in the IUCN Red List – Critically Endangered, Endangered and Vulnerable. The definition of each level is set out on this page from BirdLife.

This flock of Far Eastern Curlew has just been spooked. Disturbance is a major issue at some key sites. Julie Keating won Environmental Citizen of the Year for her work in supporting shorebirds in Port Hacking (NSW)

Several of the wader species on the East Asian-Australasian Flyway that are most at risk, such as Nordmann’s (or Spotted) Greenshank and Spoon-billed Sandpiper, are not included in the review by Hansen et al. because they do not generally migrate as far south as Australia (but there is a WaderTales blog called Spoon-billed Sandpiper: track and trace).

Two species that winter in Australia in significant numbers are categorized as Endangered by IUCN – Far Eastern Curlew and Great Knot. When comparing the Hansen estimates (mainly based on data from 2011 to 2016) with Bamford et al (data from 1986 to 2000), the population of Far Eastern Curlew is now assessed to be lower (35,000, as opposed to 38,000), while the population of Great Knot is now thought to be bigger than previously realised (425,000, as opposed to 375,000). This does not mean that numbers have gone up – we know that annual counts at key sites have fallen significantly – it indicates a more comprehensive estimate of numbers, particularly in uncounted areas.

Indeed, as shown by Colin Studds et al, summarised in Wader declines in the Yellow Sea, both Far Eastern Curlew and Great Knot numbers have been declining at an alarming rate for two decades. The fact that the drop in Far Eastern Curlew numbers is less striking than might have been expected and that there is an apparent increase in Great Knot numbers reflects more comprehensive counts from what are now known to be important sites along the flyway. The increased figures do not mean that the two species are any less threatened!

Seven species considered in the Hansen paper are listed as Near Threatened. Estimates of Grey-tailed Tattler and Red-necked Stint have been raised, from 50,000 to 70,000 for the tattler and from 325,000 to 475,000 for the stint. There is no change in the estimate of Bar-tailed Godwits (325,000), despite significant declines in numbers in Australia, again reflecting the discovery of significant numbers in sites where they were previously unrecorded or under-recorded. The Black-tailed Godwit estimate remains at 160,000.

The estimates of two Near Threatened species have dropped alarmingly, despite more comprehensive information on the sites that they use. Red Knot and Curlew Sandpiper numbers have halved (220,000 down to 110,000 and 180,000 down to 90,000, respectively).

Cautionary tale

The figures in the Hansen paper are based on the best available information for the period up until 2016. The welcome, increased focus upon shorebird conservation issues throughout the flyway is uncovering new and better count data. Birgita shared the following cautionary tale about the Near Threatened Asian Dowitcher, the population estimate of which is given as 14,000 in the paper, somewhat lower than the estimate of 24,000 that was published by Bamford et al in 2008.

“In 2019 there was a count of 22,432 Asian Dowitchers at Jiangsu Lianyungang, on the Chinese coast, which was interpreted as being 97.5% of the global population at the time (based on Bamford). This figure well exceeded our 2016 estimate of 14,000, clearly indicating that, despite access to greater volumes of data through the Asian Waterbird Census, there was a substantial number of dowitchers that had been missed in standard monitoring. So we can see already how, even with the extrapolations we used, the numbers can be highly uncertain.”

Population estimates are only estimates – the clue is in the name – but it is important to work out, as best as possible, how many birds there are, in order to assess the vulnerability of individual species and the importance of the sites that are used by assemblages of shorebirds. When 1110 Curlew Sandpipers were counted at Broome (Western Australia) in 2021, that was over 1% of the reduced flyway total of just 90,000 – and that’s important. Similarly, 4674 Bar-tailed Godwits in Tasman Bay (New Zealand) in 2021 exceeds the 1% threshold for the species and an amazing count of 9810 Sharp-tailed Sandpipers at Lake Martin (Victoria) in 2019 represents 11.5% of the 85,000 estimate in the Hansen paper. There are many more examples.

This population information is complementary to regular counts at key sites, that track population trends, and colour-ring observations that monitor annual survival rates of adults (there is a WaderTales blog about this – Measuring Shorebird Survival).

Shorebirds at Jiangsu, on the Yellow Sea coast of China – now a conservation area

More details

In the Ibis paper, Birgita Hansen and her colleagues discuss the rationale and limitations of the approaches they have used to obtain population estimates and how their methods could be used in other situations. Data available for population estimates will always vary in quality and extent among species, regions and migration stage, and approaches need to be flexible enough to provide relevant information for conservation policy and planning. Anyone considering this sort of exercise would be advised to read the whole paper:

Generating population estimates for migratory shorebird species in the world’s largest flyway. Hansen, B.D., Rogers, D.I., Watkins, D., Weller, D.R., Clemens, R.S., Newman, M., Woehler, E.J., Mundkur, T. and Fuller, R.A. Ibis. DOI/10.1111/ibi.13042


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.

New Bar-tailed Godwit Subspecies

The description of a new subspecies of Bar-tailed Godwit begs two questions, ‘How do we identify new subspecies?’ and ‘Is the concept of a subspecies helpful?’. Put simply, the answers are ‘That depends’ and ‘Yes, especially when subspecies are the focus for conservation action’.

A new study of Bar-tailed Godwits, and the proposal to identify Limosa lapponica yamalensis as a new race, usefully highlight the importance of the flyway between Siberia and the Arabian Sea and the challenges being faced by waders that use this migration route.

Limosa lapponica yamalensis

Bar-tailed Godwits breed across the whole of northern Eurasia and in Alaska. Since the last Ice Age, following glacial retreat, new breeding opportunities in northern latitudes have become available to waders. The emergence of separate migration flyways, linking wintering and breeding areas, has led to a divergence of the species into a number of distinct populations, some of which have already been defined as subspecies. In their paper in IBIS, Roeland Bom (NIOZ Royal Netherlands Institute for Sea Research) and colleagues from the Global Flyway Network argue that the taymyrensis subspecies should be further divided between those that winter in the Middle East (and presumably East and South Africa) and those that winter in West Africa. Their recommendation is based on studies of migratory behaviour, breeding area, morphology (measurements) and population genetic differentation in mitochondrial DNA. Their main focus has been a study of spatial and temporal differences in migration routes.

How many subspecies?

The most famous subspecies of Bar-tailed Godwit is baueri; these amazing birds fly directly from Alaska to New Zealand at the end of the breeding season and return via the Yellow Sea in spring. Theirs is the most impressive migration route of any wader, including non-stop flights of well over a week. These are the largest Bar-tailed Godwits.

Bar-tailed Godwits wintering in Australia are mainly birds of the menzbieri race. These breed from central Siberia across to north-east Russia and migrate north and south via the Asian coast, especially the Yellow Sea. Bar-tailed Godwits breeding further south in eastern Russia are usually described as anadyrensis, these birds are smaller than baueri but bigger than menzbieri.

Bar-tailed Godwits breeding in western Siberia have all been known as taymyrensis, named after the Taymyr** Peninsula, while those breeding in Scandinavia, Finland and the far west of Russia are identified as lapponica. Nominate Limosa lapponica lapponica birds winter in northwest Europe and most taymyrensis use the East Atlantic Flyway to winter at least as far south as Guinea-Bissau in West Africa. Taymyrensis is the smallest race. Lapponica and taymyrensis were recognised as separate subspecies two decades ago and this differentiation has been useful in discussions about Bar-tailed Godwit conservation. In the UK we see both lapponica and taymyrensis, as discussed in Bar-tailed Godwits: migration and survival.

Until now, birds migrating south from western Siberia, as far as East and South Africa, have been considered to be taymyrensis and it is these birds that have been reclassified – with the new name of yamalensis, after the region of Russia in which the subspecies breeds.

** In the IBIS paper, the authors use Taimyr (rather than Taymyr) as this is preferred by Russian co-authors. I am maintaining consistency within WaderTales.

What’s different?

The studies summarised in the IBIS paper by Bom et al are designed to test whether Pavel Tomkovich, who has studied Russia’s artic-breeding waders for several decades, was right when he suggested that taymyrensis should be split into two subspecies. (See Population structure and migratory links of Bar-tailed Godwits: Current knowledge and unsolved issues in Achievements in Studies on Waders of Northern Eurasia). Based on a small number of movements of ringed birds between areas where there are few birdwatchers, Pavel suggested that L. l. taymyrensis likely comprises two distinct (flyway) populations, one wintering in the Middle East, West Asia and East Africa and breeding on the northern West-Siberian Plain, and the other wintering in West Africa and breeding on and around the Taymyr Peninsula.

Using satellite-tracking, Roeland Bom and colleagues have now described the migration routes, breeding destinations, and annual-cycle timing of Bar-tailed Godwits using wintering areas in the Middle East (Oman) and West Africa (Mauritania and Guinea-Bissau). To understand further the extent to which the two groups are different and whether or not there is any mixing of the two populations, they also examined differences in mitochondrial DNA.

Tracking through space and time

Members of the research team attached solar-powered tags to eastern taymyrensis Bar-tailed Godwits in Oman and western birds in Mauritania, Guinea-Bissau and the Wadden Sea. 52 birds caught between 2015 and 2018 provided usable tracks; 11 birds (12 tracks) linked the western wintering/migration route to the area in and around the Taymyr peninsula, while 9 birds (19 tracks) linked the Middle East to the northern part of the West Siberian plain. To limit potential negative effects of carrying extra weight, only the largest birds were tagged, which means that all of the tags were deployed on females.

Bar-tailed Godwits from wintering areas in the Middle East staged for several days in the areas around the Caspian and Aral Seas, during both northward and southward migration. Both of these areas were already thought to be important for passage waders but analysis of tracking data emphasise the critical role they play in the annual cycle of Bar-tailed Godwits. Some birds also spent short periods of time at sites in the United Arabian Emirates, Iran and India. After leaving their breeding sites, some birds moved up to 1,000 km north, to feed in high-Arctic coastal Siberia, before embarking on southward migration.

Bar-tailed Godwits tracked from wintering sites in West Africa staged in the Wadden Sea, during both northward and southward migration, with other staging points in Spain, Portugal and France, during northward migration. From the Wadden Sea, most West Africa birds flew directly to the northern West-Siberian Plain (near or in the breeding area of the Middle East birds) before heading for the Taymyr Peninsula. All birds routinely moved north before leaving for the Wadden Sea, with many using the same fuelling areas as Middle East birds.

Although birds from West Africa and the Middle East used the same feeding areas in spring and post-breeding, the phenology of the two populations were different. The northward migration, the arrival in the staging and breeding sites and the southward migration of Middle East birds were earlier than for the West Africa birds. Some birds of the two groups could be found in the same areas in Siberia in the pre- and post-breeding staging, emphasising the fact that timing differences can be just as important as spatial separation when it comes to the evolution of subspecies. This ‘spatial overlap but with timing differences’ story is similar to that seen in limosa and islandica Black-tailed Godwits in The UK and The Netherlands, as discussed in Godwits in, godwits out: springtime on the Washes.

Body size and shape

The measurements of taymyrensis and yamalensis overlap but there is a tendency for Middle East birds to have smaller bills and longer wings than West African birds. The authors suggest that, given that the two subspecies have similar diets during the non-breeding season, the different bill structure may be related to feeding requirements during the breeding season, with taymyrensis breeding in open tundra and yamalensis breeding in forest tundra and bogs within the boreal zone. It is interesting to note that the morphological differences between the taymyrensis and yamalensis subspecies are greater than those between taymyrensis and lapponica.

Genetics

West African and Middle East birds could not be separated using genetic tools available to the researchers. The authors point out that there is very little genetic variation between the three more western subspecies (lapponica, yamalensis and taymyrensis), indicating that either these three populations may have diverged recently (i.e. well after the last Ice Age) or that there is still some (small) geneflow between subspecies. The authors dicusss this in depth, in the context of all known populations of Bar-tailed Godwits and of waders in general.

A new subspecies

Roeland Bom and colleagues conclude that the old taymyrensis taxon consists of two distinct populations with mostly non-overlapping flyways, which warrant treatment as separate taxonomic units. They argue that ‘separation in space and time’ can define separate subspecies and that these differences will become apparent before morphological (size and shape) differences develop and long before it will be possible to spot genetic differences.

Breeding locations of tagged Bar-tailed Godwits

Yamalensis Bar-tailed Godwits breed on the northern West-Siberian Plain including the Yamal Peninsula. Birds of the subspecies follow the Central Asian Flyway, with main stopover sites in the Caspian Sea and the Aral Sea. Satellite tracking has connected Oman with other wintering areas in the Middle East, Iran, Pakistan and West India. It is likely that some birds continue further south on the West Asian – East African Flyway, with two, earlier ring recoveries showing that the subspecies can winter as far south as South Africa.

Conservation considerations

One of the key benefits of defining yamalensis as a new subspecies is that priority setting (and hence funding) for conservation action is often defined at the subspecies level. As we learnt from Ashwin Wisvanathan and Les Underhill at the International Wader Study Group conference in 2021, the numbers of waders spending the non-breeding season at the southern extremes of the Central Asian Flyway (in India) and of the West Asian – East African Flyway (in South Africa) have declined alarmingly over the last few years.

Roeland Bom and colleagues quote population estimates for taymyrensis and yamalensis Bar-tailed Godwits of 600,000 and 100,000-150,000 respectively but these are old numbers and West African winter populations are known to be declining. It is important to establish new population estimates and trends for the two populations and better to understand the migration of yamalensis. Potentially, more satellite tracking might help to suggest where to look for wintering yamalensis along the vast coastline from the southern tip of Africa to the southern tip of India, and to identify further spring and autumn stop-over sites.

Paper

The paper in IBIS concludes with a detailed description of the subspecies Limosa lapponica yamalensis, together with measurements that separate it from other races. You can read more in:

Central-West Siberian-breeding Bar-tailed Godwits (Limosa lapponica) segregate in two morphologically distinct flyway populations. Roeland A. Bom, Jesse R. Conklin, Yvonne I. Verkuil, José A. Alves, Jimmy de Fouw, Anne Dekinga, Chris J. Hassell, Raymond H.G. Klaassen, Andy Y. Kwarteng, Eldar Rakhimberdiev, Afonso Rocha, Job ten Horn, T. Lee Tibbitts, Pavel S. Tomkovich, Reginald Victor & Theunis Piersma. IBIS.

Other WaderTales blogs about waders on the West Asian – East African flyway

Well-travelled Ring Plovers makes a link between North Africa and the furthest northeast corner of Russia, extending the reach of the West Asian – East African Flyway further east than sometimes shown on maps.

In search of Steppe Whimbrel describes the migration of Numenius phaeopus alboaxilliaris, a Whimbrel subspecies that has already been declared extinct once. The subspecies migrates between the steppes of Kazakhstan and Russia and coastal East Africa.

Following Sociable Lapwings describes the migration of this threatened species along both the West Asian – East African Flyway and the Central Asian Flyway, with some interesting thoughts about a migratory divide in a wader which exhibits relatively low philopatry.


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.

The flock now departing

“The flock now departing from the tideline is bound for Beauvais. Curlew can change here for destinations in Germany and Russia”. It’s fascinating to wonder what might be happening when a flock of waders takes to the air, gains height and sets off in a particular migratory direction. With more individuals wearing tracking devices, it was only a matter of time until someone would have data that provides clues as to the association of individuals within flocks – as we see in a 2021 paper in Bird Study by Frédéric Jiguet and colleagues: Joint flight bouts but short-term association in migrating Eurasian Curlews.

Setting off on migration

When we get on a plane to a particular destination, everyone else who is on the same journey has chosen to travel at the same time and we all know where we are going. Each of us has checked that we have what we need for the journey and has a plan of what to do when we land – whether that involves a short shuttle to home or a lay-over before catching another flight.

For waders, planning must be more random? It’s presumably safer and more efficient to be part of a flock but how do you know which flock to join, who organises the schedule and is information shared? We can get some clues from observations of departing migratory flocks. In estuaries, there is often the chatter (which is hard to interpret but tells us that something is about to happen), then the first birds take to the air and start to gain height. A few birds may peel off and return to the tide-line while other birds take off and catch up with the departing flock. As the birds gain height, the direction of travel becomes clearer and more birds may decide to return to the mudflats. There is now a migratory flock of birds that are committed to flying in a particular direction. We have no idea how that direction was chosen, of course, but there is a plausible explanation as to how the flock might have formed.

This is not the last decision that members in a flock might need to make. Tired birds may need to drop out of the flock, to take a break. Perhaps some birds might realise that the direction of travel does not work for them and the flock might break up?

It can be just as chaotic when a flock reaches a destination. Watching Black-tailed Godwits arriving in South Iceland in April is fascinating; a tired flock might come in off the sea, land and start drinking, before either resting or feeding, but this is not always the case. On a clear day with fair winds, the flock may split up, with some birds keen to keep flying and others happy to stop. This reinforces the impression that a flock only maintains its integrity as long as being in a group meets the needs of the individuals it contains.

Tracking Eurasian Curlew

Understanding migration is an important element of Curlew conservation studies in France.

In their Bird Study paper, Frédéric Jiguet and colleagues describe four cases of joint migration by tagged Eurasian Curlews. Their observations were a biproduct of research aimed at a better understanding of the origins and migration patterns of Curlew that spend the winter in France. The species has been a popular target for French hunters, many of whom are keen to resume shooting, as you can read in the WaderTales blog Black-tailed Godwit and Curlew in France. It is estimated that more than 7000 Curlew were shot in France annually prior to 2008, when the first moratorium was put in place.

There is an urgent need to understand links between wintering sites and breeding sites, especially in areas where the species is in rapid decline. How important is France to the Curlew that breed in countries such as Poland and Germany? The current ban on shooting is not perfect (see paper in Forensic Science International: Animals and Environments) but it is better than nothing, given rapid declines in Curlew numbers across Europe.

In winter and spring 2020, the research team deployed 61 GPS tags on Curlews in France and Germany, hoping to learn more about breeding ecology and migratory connectivity. In a separate study, in Poland, four captive-bred juvenile curlews were tagged and released in July 2020. Between them, these tagged birds led to four cases of joint migration bouts. One case concerned two adults leaving their wintering ground for the pre-breeding migration. Two other cases were birds leaving their breeding grounds at the start of migration. The last one was of two juveniles initiating their first flights to the non-breeding grounds.

Spring migration

About 27,500 Curlew spend the winter in France (see French report produced jointly by government and shooting groups), representing about 5% of the European population. Tracking has shown that these birds breed in Belgium, Germany, Sweden, Finland, Austria and Russia (see article published by Bird Guides) but there are reports of ringed birds from many other countries, including the threatened populations in Poland and the UK.

Thousands of Curlew spend most of the year in coastal France – representing 5% of the European population.

Return migration to breeding areas takes place in early April. Frédéric Jiguet reports ‘groups of curlews rising high in the sky at sunset’ from the Moëze-Oléron and Baie de l’Aiguillon Nature Reserves in southwest France.

Back-mounted GPS tag

On 17 April 2020, two individuals wearing tracking devices left their French wintering site at sunset, between 22:37 and 22:40. They became closely associated just ten minutes prior to the start of migration, having typically stood 100 metres apart during the previous hour. They flew together for seven hours before making a stop-over north of Paris, between Creil and Beauvais, in the Thérain Valley.

  • 200185 was on its way again two hours later, flew for six hours, stopped again in the Netherlands and arrived in Norderney, an island of the Wadden Sea in northern Germany, at 18:39 on 19 April.
  • 200187 had a much longer layover in the Thérain Valley, making another evening departure at 20:05 on 18 April. It continued migrating, in stages, for more than a month, crossing the Ural Mountains and reaching the Yamalia municipality, in Asian Russia.

Two birds that had been on the same flight from southwest France ended up in very different locations and at very different times. The German Curlew reached its summer destination five weeks before the Russian bird arrived on territory, the latter having secured places on several different ‘international flights’ as it made its way east and north (see figure below).

Post-breeding migration

After breeding, adult Curlew head towards wintering sites, perhaps stopping to moult en route. Some birds do not travel far; for instance, there are colour-marked birds that winter on the Wash (eastern England) and fly just a few kilometres inland to breed. The Bird Study paper includes reports of two occasions when tagged birds have been spotted migrating together from German and French study areas. Southerly migration of all four birds commenced during the evening of 17 June 2020.

French birds: Two individuals departed simultaneously from Deux-Sèvres (central France) between 19:16 and 19:17 for a non-stop southward flight and arrived together at Ria de Treto estuary, in northern Spain on 18 June at 05:49. The two birds departed separately from this stopover site the same day (18 June).

  • 200201 departed at 18:18, for a non-stop flight to Kenitra (Morocco) where it stopped briefly, before moving a short distance north to Merja Zerga.
  • 200204 departed at 19:46 and flew to the Atlantic coast of Spain, stopping for 2.5 hours on Isla Cristina and then flying to its final destination at Ilha de Tavira, in southern Portugal.

After separation, the two birds travelled at different times but followed quite similar routes and even flew at similar altitudes.

German birds: On 17 June 2020, two individuals departed simultaneously from Dollar Bay, in the Wadden Sea National Park. 201075 began migration between 18:58 and 19:03. After five kilometres, if flew over 201072 at an altitude of about 190m. The latter bird took off and joined 201075. They then flew together for five hours, landing in the Rhine-Meuse-Delta (Netherlands).

201075 departed from the Rhine-Meuse-Delta on 20 June and, after one more stop-over, reached its final destination on the Brittany coast on the evening of the next day.

201072 was also bound for Brittany. It departed on 23 June and flew non-stop for six and a quarter hours.

Migration of juveniles

It will be hard to satellite-tag enough wild juvenile waders to pick up instances of marked individuals migrating in the same flocks. However, head-starting may give some clues as to what might happen when naïve flocks of juvenile waders start their migratory journeys, months after the parents have left them. The full story is told in the paper but a quick summary tells us that two Polish head-started Curlews were released on 1 July, departed together on 5 August and landed in the Baie de l’Aiguillon (France) on 8 August. In between times, they came close to landing in The Netherlands, flew along the English coast from Dover to Poole, flew a long way south and west around the Bay of Biscay and then northeast to the coast of France. They both spent the winter in the Baie de l’Aiguillon but not together.

Although it will be difficult to compare the migratory behaviour of wild-caught and head-started wader chicks using satellite tags, just because of probabilities and costs, researchers are building up datasets using smaller geolocators and GPS tags. Here’s hoping that we will soon know more.

Paper

The nutrient-rich mud of Ile Madame

This paper provides observations of just four instances of joint migration but each story is fascinating. They give us insights as to what might be possible as devices get smaller and when land-based tracking stations collect signals from passing birds. For the moment we can use our imagination to interpret the chattering of pre-migratory flocks of waders, the appearance of a small flock of waders at an inland spot in spring and the noisy arrival of a lone Curlew on an estuary in June.

The paper contains a lot more detail about the methods used to collect and interpret data and a discussion that sets Curlew migration within a much broader conceptual context. Here’s a link:

Joint flight bouts but short-term association in migrating Eurasian curlews.

Frédéric Jiguet, Pierrick Bocher, Helmut Kruckenberg, Steffen Kämpfer, Etienne Debenest, Romain Lorrillière, Pierre Rousseau, Maciej Szajdaand & Heinz Düttmann. Bird Study. DOI/10.1080/00063657.2021.1962805

Wintering Curlew from as far away as Russia and Sweden can be found roosting in these French saltmarshes

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.

Flying high with Great Snipe

As tagging devices get smaller and more sophisticated, they are revealing even more wonders of shorebird migration. We already know that Great Snipe are amazing – flying up to 7,000 km non-stop – but a 2021 paper by Åke Lindström and colleagues describes a striking daily cycle of altitude change during their long migratory journeys.

In their new paper in Current Biology, Åke Lindström and colleagues have used activity and air pressure data from multisensor dataloggers to show that Great Snipes repeatedly changed altitudes around dawn and dusk, between average cruising heights about 2,000 m (above sea level) at night and around 4,000 m during daytime. Most birds regularly flew at 6,000 m and one bird reached 8,700 m, an altitude that is just 150m short of clearing the top of Mount Everest! The same daily cycle was apparent everywhere – independently of climate zone, habitat and the height of the land being overflown. Wherever they are, as morning breaks migrating Great Snipes gain altitude – but why?  

Great Snipe

Great Snipe spend the winter in Africa, between 20 degrees south and 15 degrees north of the equator, heading north in spring to breeding areas in Scandinavia and northern Europe, as far east as 95°E (which is the same longitude as Myanmar).

The first paper to reveal the remarkable migration of Swedish Great Snipe appeared in Biology Letters in 2011, when Raymond Klaassen, Åke Lindström and colleagues revealed the tracks from three birds carrying geolocators. During these journeys, these individuals made long and fast autumn movements, covering between 4300 and 6800 km in two to four days and overflying suitable stopover sites that were used in spring. Ground speeds of 15 to 27 m/s are equivalent to between 54 and 97 km/h (33 to 60 miles per hour).

In 2016, using 19 tracks from four years of captures and recaptures, the same team published a paper in the Journal of Avian Biology. About half of the birds flew directly from the breeding grounds to sub-Saharan Africa, the others making a few shorter flights down through Scandinavia before embarking on a long trans-Sahara flight. Birds took advantage of wet seasonal conditions in this Sahel region for three weeks before moving south to the Congo Basin. Spring migration consists of a rapid movement across the Sahara, only a little shorter than the very long non-stop autumn fights, followed by slower movements through Eastern Europe. Birds arrived back on breeding areas in mid-May.

Great Snipes spend about eight months of the year in sub-Saharan Africa. In a 2017 paper in Wader Study, Edouard Debayle, Åke Lindström and colleagues analysed the moult and fattening patterns of over 3,000 hunted birds, to try to learn more about the phenology of migration. They discovered that:

  • Adult males arrived in Africa from mid-August, having started and suspended the moult of the main flight feathers before arrival.
  • Females on average arrived somewhat later and were about one month behind in the progress of flight feather moult.
  • The adults of both sexes resumed primary and secondary moult immediately upon arrival and typically completed it by the end of November, in males, and the end of December in females. Juvenile Great Snipes arrived later than adults and did not moult their flight feathers in the first autumn/winter.
  • Males apparently departed northwards between late March and late April, and the females about two weeks later. There is information about rates of fat deposition in the paper.
Great Snipe watches on as team members set a mist net

Flying high

Processing the catch

Several factors could influence the flight altitude of migratory birds. For example, how high is the land over which birds fly, what are the best temperatures and humidity conditions for efficient flight, at what height can a bird find the most helpful winds, can a bird use navigation landmarks and how can predation best be avoided?

Recent tracking of migratory birds of a range of species has shown that individuals change flight altitude more commonly and dramatically than previously thought but why? In their paper in Current Biology, Åke Lindström and colleagues reveal information from 25 tracked Great Snipe journeys and discuss the reasons that may lie between the patterns that they see.

Methods

Information about behaviour and flights was collected from multisensor dataloggers, consisting of an accelerometer for activity measurements, a barometric pressure sensor with internal temperature sensor, a light-level sensor, a real-time clock, and memory. The dataloggers weighed 1.4–1.7 g (about 1 % of a bird’s total body mass) and were attached to a plastic ring on the bird’s tibia.

In total, 107 dataloggers were put on Great Snipes between 2015 and 2019. Of these, 36 birds were retrapped one, two or three years later (but four birds had lost their loggers). This is an overall recapture rate of 34%, which is similar to the figure for ringed birds. In total, 25 out of the 32 retrieved loggers had functioned for some time, and 16 carried information on flight altitude for at least one of the long flights.

The methods section provides full details of how geolocator data were interpreted and altitudes were calculated and adjusted. Information on air temperatures at different altitudes and the topological features on flight paths provided a background against which to try to understand migration patterns.

How high?

The new data from a small number of multisensor data-loggers greatly enriched the migration story of Swedish Great Snipe, as revealed byÅke Lindström and colleagues. The key results in the Current Biology paper are:

Breeding habitat in Sweden

Flight duration: There were three long flights, two legs on the way south and one on the way north. As noted earlier, northerly migration slowed once birds landed in Europe.

  • On average, male Great Snipe left Sweden on 24th August and flew across Europe and the Sahara for 73.4 hours, before landing in the Sahel.
  • The mean departure date from the Sahel was 24th September. An average of 23.2 hours later a bird would land in the Congo basin.
  • Northerly flights commenced on 18th April, lasting an average of 82.4 hours and concluding in Europe.

Cyclic flight altitudes: There was an overall strong and consistent daily cycle in the altitudes used by the Great Snipes, in all three long flights. After a night at moderate to high altitudes the birds ascended to very high altitudes in early morning, stayed at these levels during the day, and descended again in late afternoon. They then repeated this cycle for one or two more days.

  • The mean individual daytime flight altitude in the first Autumn flight was on average 4,549 m, compared to 2,126 m at night.
  • For the In-Africa autumn flights, comparative figures were 3,874 m in daytime and 1,860 m at night.
  • For the Spring flights, comparative figures were 4,114 m in daytime and 1,612 m at night.

These altitudes were estimated from air pressure readings and may be underestimates. For comparison, the highest point in the Alps is 4807 m.

Peak altitudes: Some Great Snipes occasionally flew extremely high and then always during daytime. Three birds in Autumn and two birds in Spring reached 7,000 m or more. Migratory waders are able to carry out flapping flight at such high altitudes due to several physiological adaptations of the heart, lungs and muscles. The single highest altitude estimate of 8,077 m was reached in Autumn. If air pressure is accounted for this bird may have been flying above 8,000 m for five hours, perhaps reaching an altitude of 8,700 m, and coping with an air temperature of -21.3 °C. Putting this in a local context for readers: Mount Everest is 8848m high and the high points in other regions of the world are: Africa (5895 m Kilimanjaro), South America Aconcagua 6959 m, North America Denali 6190 m and New Zealand Mt Cook 3754 m.

Ambient temperature, wind condition and humidity appeared not to influence the differences in day and night altitudes chosen by Great Snipes.

Discussing the results

Great Snipe with a datalogger

The daily pattern of altitude changes for Great Snipes was very similar between Autumn, In-Africa and Spring flights, suggesting a common cause that is largely independent of climate zone (temperate or tropical), topography and landscape overflown (forest, savanna, farmland, desert or water). Altitude changes have been reported for other waders, such as Black-tailed Godwits, that have been linked to both ambient temperatures and finding more beneficial winds.

The authors discuss the possibility that landmarks are easier to see from a higher altitude when flying in daylight and suggest that predator avoidance may also account for higher day-time elevations. It would be interesting to know if a daily cycle is apparent in long flights over areas largely lacking bird predators, such as vast oceans.

There is no daily cycle in ambient air temperature or wind conditions at high altitudes that could explain the overall regular pattern of flight altitude selection found in Great Snipes but the authors discuss the theory that the warming effect of solar radiation may be countered by flying through colder, higher air. The temperatures at these heights would be too cold at night.

There are still few papers that provide altitude data for long-distance migrating birds but all of them report altitude changes and have revealed some surprisingly high flight altitudes. With more studies we may well find that migration is even more impressive than we already thought!

Paper

The full methods, results and discussion can be read in the paper in Current Biology.

Extreme altitude changes between night and day during marathon flights of Great Snipes Gallinago media: Åke Lindström, Thomas Alerstam, Arne Andersson, Johan Bäckman, Peter Bahlenberg, Roeland Bom, Robert Ekblom, Raymond H. G. Klaassen, Michał Korniluk, Sissel Sjöberg & Julia K. M. Weber.


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.

Dunlin: tales from the Baltic

Veli-Matti Pakanen, Kari Koivula  and colleagues have been studying Finnish Dunlin for nearly twenty years. These are schinzii birds that breed in coastal grazing meadows around the Baltic Sea. Several papers have been published, as you will see below, some of which are based on information collected using geolocators attached to leg-flags. Dunlin numbers in coastal Finland are dropping quickly, so this research is important to the conservation of the species. In a 2020 paper, Veli-Matti and colleagues ask whether intensive geolocator-based studies are having a negative effect on individual birds in this already-declining study population.

Global Dunlin

Up to ten races of Dunlin have been identified, which together encircle the globe. Most Dunlin spend the non-breeding season in the northern hemisphere and all migrate north in spring. The breeding and wintering ranges of the various races are summarised at the end of this blog. Here, the focus is on Baltic schinzii Dunlin, a small part of a race that largely winters in coastal North Africa and breeds in southern Scandinavia, Northern Europe, the British Isles, Iceland and southern Greenland.

According to Wetlands International, there were between 4.3 and 6.8 million Dunlin worldwide in 2015, with about one million breeding birds within Europe (BirdLife International). Whilst it is acknowledged that numbers are declining, the large range of the species means that it is still considered to be of ‘least concern’.

Different populations are faring differently. In his description of the changing Dunlin distribution in European Breeding Bird Atlas 2, John Calladine points out that there have been major losses across Europe, including Britain & Ireland, and that Baltic populations ‘are now considered threatened’. The Baltic schinzii population was most recently estimated as between 500 and 640 pairs – less than a fifth of the estimate in the 1980s. John highlights gaps in knowledge that research by the team in Finland and other groups elsewhere are helping to fill.

Migration and survival of schinzii

Dunlin that visit the UK are mostly of the schinzii and alpina races, as indicated in this map. There has been a noticable drop in the number of schinzii birds stopping off in July.

In pre-geolocator days, Ole Thorup and colleagues analysed recovery information available for Dunlin breeding around the Baltic, using 40 years of ringing data from Finland, Sweden, Denmark and Germany. At that stage there were only six mid-winter recoveries in N and NW Africa. The analysis emphasised the importance of wintering and stop-over sites in the Baltic, the Wadden Sea, SE and S England, the Atlantic seaboard of France, and the Iberian Peninsula. Paper in Ardea (2009).

Pakanen et al investigated changes in the survival rates of schinzii Dunlin nesting in Denmark, based on ringing and recaptures of a marked population. They found that annual apparent survival rates dropped from 0.817 to 0.650 between 1990 and 2006, equivalent to a doubling of the chance of dying in any given year. Paper in Bird Study (2016). The importance of monitoring survival rates is discussed in the WaderTales blog: Measuring shorebird survival.

Nests in these flat coastal marshes along the Baltic coast are increasingly susceptible to summer flooding

The use of geolocators enabled Veli-Matti Pakanen to add more detail to the migration story in the 2018 paper, Migration strategies of the Baltic dunlin: rapid jump migration in the autumn but slower skipping type spring migration. He and his colleagues showed that autumn migration is faster than spring migration, characterised by fewer stationary periods, shorter total stopping time and faster flight. The Wadden Sea was found to be an important autumn staging area for all of the tagged birds. Some birds stopped once more before reaching Mauritania. On spring migration, more sites were visited on the way north. The important conservation message from the paper is that Baltic Dunlin may be especially vulnerable to rapid environmental changes at their staging and wintering areas. (In Travel advice for Sanderling there is a suggestion that annual survival is relatively low for birds that winter in Mauritania).

In a 2015 paper in Ornis Fennica, Pakanen et al reported on the results of a single-year analysis of survival rates, concluding that there were no strong effects of leg-flagged geolocators on return rates or reproduction in schinzii Dunlin. However, they did suggest that “long term evaluations that capture the full suite of environmental conditions and assess impact on brood care are needed”. This is a question which leads us neatly on to the 2020 paper: Survival probability in a small shorebird decreases with the time an individual carries a tracking device.

Long-term tracking of individuals

To understand the range of pressures that migratory birds face, one needs to know where individuals spend their time, as was demonstrated in Spoon-billed Sandpiper: Track & Trace and Teenage Waders. Waders of a range of species have been wearing geolocators for ten or more years now, either the same devices on birds which have evaded capture, or a series of tags, when data have been downloaded and replacement tags fitted. These long-term surveys are developing our understanding of the repeatability of migratory behaviour and how birds deal with variable weather patterns, but is there a long-term cost to the individual birds that are tasked with finding out this important information?

This Dunlin wore a ring-mounted geolocator for one year, to collect movement data

Effects of tracking devices on survival are generally considered to be small. However, most studies to date have focused on birds that were caught in one breeding season and recaptured in the following season, to retrieve the geolocator and download the data. In their 2020 paper, Veli-Matti Pakanen and colleagues were able to investigate the possible accumulation of negative effects when individuals have carried the tracking devices for longer periods. Survival rates for tagged birds were compared with 338 colour-ringed birds that were followed for all or part of the period 2002 to 2018.

In the summers of 2013 & 2014, fifty-three adult schinzii Dunlin were fitted with leg-flag mounted geolocators, with a mass equivalent to 1.5 – 2.0% of body-mass. Of these birds, 17 tags were retrieved after one year and 9 after two years. Other marked birds could not be caught and carried their tags for longer periods (3 or 4 years). The research team found that Dunlin carrying a geolocator had reduced chances of survival. Their models suggest that annual survival of colour-ringed males was 0.813. For a bird that carried a geolocator for a year, annual survival probability declined to 0.748 and to 0.581 for birds that carried the geolocator for at least 2 years. Their data suggest that the reduction in survival rates was greater for females than males, even though females are larger than males.

Summer flooding of coastal breeding area is becoming an increasing problem, and likely to get worse with sea-level rise and more chaotic weather patterns

In a thorough Discussion, the authors consider reasons why tags on small waders may be reducing survival, either through ongoing stresses, impacting on things such as feeding efficiency and the energy needed during migration, or because the extra burden means that tagged birds find it harder to cope with occasional periods of tough environmental conditions. They comment on the condition of the skin under removed geolocators – something that other researchers night want to look out for.

As anyone studying breeding waders will know, nest-trapping to retrieve tags is not easy, especially if adults lose their clutches when incubation has only just started, due to flooding, predation etc. Birds may end up carrying tags for longer than intended. The authors “recommend that the detrimental effects of tagging may be avoided by developing attachment methods that are automatically released after one year, e.g. biodegradable materials”.

Balancing costs and benefits

The results from the Pakanen study of long-term survival suggest that requiring a small wader to carry a geolocator for several years may have an impact on survival. As in all mark-recapture studies, researchers are urged to assess the costs to the individual when seeking to understand what might be affecting the viability of a population.

Four previous WaderTales blogs have discussed tag effects:

Details of the Dunlin tagging effects study:

Survival probability in a small shorebird decreases with the time an individual carries a tracking device.

Veli-Matti Pakanen, Nelli Rönkä, Thomson Robert Leslie, Donald Blomqvist, Kari Koivula. Journal of Avian Biology (2020): https://doi.org/10.1111/jav.02555

Four other papers relating to this Finnish Dunlin study

Grazing by cattle is an important management tool in coastal meadows. Pakanen et al studied the impact of trampling on artificial nests and concluded that even recommended stocking rates were too high for chick numbers that could deliver a sustainable population. Paper in Biodiversity and Conservation (2011). (Redshank on British estuaries are similarly vulnerable to trampling – see Big Foot and the Redshank Nest).  

In a follow-up paper in 2016, Pakanen et al concluded that Dunlin populations could be sustained in grazed coastal meadows as long as there was no active grazing before 19 June. Meadows with grazing cattle attracted breeding birds but there was insufficient breeding success for sustainability. Paper in Ecology & Evolution (2016).

If these schinzii Dunlin chicks return to breed they are likely to try to nest nearby; something that needs to be considered when considering conservation measures

Dunlin are strongly philopatric, with both male and female chicks recruiting to suitable habitat close to natal sites. In a paper in Ibis (2017), Pakanen et al show that natal dispersal of Dunlin is strongly linked to the size of their natal site and how isolated the site is. They suggest that inbreeding may be avoided by creating a network of suitably sized patches (20–100 ha sites), no more than 20 km apart from each other. These may work as stepping stones for recruiting individuals. These results are corroborated by a 2021 microsatellite study in BMC Ecology and Evolution which shows genetic differentiation and isolation by distance within the Baltic Dunlin population.

Up to ten races

In western Europe we see three races of Dunlin – alpina, arctica and schinzii. Wintering birds are almost exclusively of the alpina race, which head north and east to northern Scandinavia, Russia and Siberia in spring. The other northern race is arctica, a very small Dunlin that breeds in low numbers in NE Greenland and possibly Spitzbergen. Schinzii has a very large breeding range, spanning the Baltic, southern Scandinavia, Northern Europe, the British Isles, Iceland and SE Greenland. There is huge variation in the timing of breeding of schinzii, as birds do not return to breeding sites in SE Greenland until the end of May, at the same time as schinzii Dunlin being studied by Veli-Matti Pakanen and colleagues in Finland will have their first young chicks.

Further east, centralis Dunlin replace alpina. Many of these birds use the Central Asian Flyway. Further east still, we find sakhalina that use the East Asian/Australasian Flyway (EAA). Two other subspecies have been identified breeding within the EAA Flyway, the more southerly kistchinski birds and actites, which breeds furthest south, on the Russian island of Sakhalin, on a similar latitude to the UK.

It is generally accepted that there are three Dunlin subspecies in North America. In autumn, arcticola head west from northern Alaska and NW Canada and follow the EAA Flyway, pacifica fly south along the Pacific coast from SW Alaska and hudsonia migrate from central northern Canada using the Atlantic Americas Flyway.


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.

Subspecies, connectivity and conservation in shorebirds

Rufa’ Red Knot in Delaware Bay

For waders such as Red Knot (Knot), conservation designations such as ‘near-threatened’ or ‘endangered’ are based upon declines and vulnerability of populations that breed in defined areas. What happens when populations mix when they are on migration or in their non-breeding areas? How do we define conservation priorities of mixed flocks? Camila Gherardi-Fuentes, Jorge Ruiz and Juan Navedo invited us to think about this issue in a 2021 Red Knot paper in Bird Conservation International.

Conservation challenges posed by overlapping subspecies

It would be convenient if subspecies of waders kept themselves to themselves but they don’t. In spring, islandica Black-tailed Godwits join limosa in Portuguese and Spanish rice fields. Icelandic populations have been increasing for a century but the Dutch population of limosa dropped by 75% between the 1970s and the period 2007-15 (as described in this blog). Which subspecies should take precedence when assigning conservation importance to a spring flock on the Tagus, or to an autumn flock in France, for that matter? These questions are not abstract; they are relevant to a decision to site a new airport for Lisbon in the estuary and to discussions about the sustainability of autumn hunting on the French coast.

Further south, in the Banc d’Arguin of Mauritania, what is the conservation importance of Dunlin? Birds from Iceland outnumber those that breed around the Baltic coast. There is no suggestion that Icelandic schinzii Dunlin are in trouble, with between 200,000 and 300,000 pairs and no indication of range change, but the Baltic schinzii population was most recently estimated as between 500 and 640 pairs – less than a fifth of the estimate in the 1980s. Does the plight of Baltic (and Irish and UK) schinzii Dunlin confer a ‘threatened’ label on the whole wintering population of the Banc d’Arguin?

Untangling Red Knot in Chile

In their paper, Insights into migratory connectivity and conservation concerns of Red Knots in the austral Pacific coast of the Americas, Camila Gherardi-Fuentes, Jorge Ruiz and Juan G Navedo present the first detailed population morphometrics of Red Knot on the southern Pacific coast of South America, during the non-breeding season, along with information about resightings of these birds throughout the Americas.

Globally, Red Knot Calidris canutus is one of the most extensively studied shorebird species and is considered as ‘Near Threatened’ at the global level (BirdLife International 2018). It is currently accepted that three subspecies are found in the Americas.

The general migratory patterns are as follows but the authors of the new paper present evidence of a more complicated picture.

  • roselaari Knot breed in Alaska and Wrangel island (Russia) and migrate along the Pacific coast to spend the non-breeding season mainly in Mexico. The total population is estimated to be 17,000 birds.
  • rufa Knot breed in northern Canada and migrate down the eastern seaboard of the Americas, some travelling as far as Tierra del Fuego. The total population is estimated to be 42,000 birds. This subspecies has been designated as ‘threatened’ in the USA, where there has been an increase in the pressure upon spring staging sites. There is a WaderTales blog about the vulnerability of this subspecies, based upon work in Delaware Bay. New genetic research is described at the end of this blog.
  • islandica Knot breed in NE Canada and Greenland and spend the winter in western Europe. Two WaderTales blogs about changing numbers of shorebirds in Great Britain and Ireland discuss declining numbers of islandica Knot.

Colour-ringing and geolocator studies that track individual birds are providing new evidence that complicates the above pattern, with some rufa Knot spending the non-breeding season on the Pacific coast of South America (Navedo, J.G. & Gutiérrez, J.S. 2019) and some roselaari wintering in Texas. Migration is even more complex, with one roselaari bird flying from Chile to Texas and then switching back west to head to Alaska (see map right). There is more about this on the Wader Study website.

The Red Knot of Chile

The team from the Bird Ecology Lab in Chile have been studying the shorebirds of the Chiloé Archipelago (42˚S, Chile) for several years. This archipelago is a Site of Hemispheric Importance for the conservation of migratory shorebirds, due to its large numbers of Hudsonian Godwit (WaderTales blog Teenage Waders) and Whimbrel.

Red Knot regularly winter in this area, with at least 150 occurring in two well-studied bays of the main island. Although it might be assumed that these birds would be roselaari, there have been colour-ring sightings of a small number of birds that had been marked with lime and green flags within the rufa flyway. With conservation of two subspecies in mind, the research team were keen to know more about the natal origins of the Chiloé Red Knot.

42 Red Knot were caught on Chiloé main island between 2017 and 2020. As well as being aged, ringed, colour-ringed with red flags, measured and weighed, blood samples were taken, in order to determine gender. The biometrics of this small sample of birds combined with sightings of red-flagged Knot has revealed a remarkable amount of information:

  • As in other Red Knot populations, males were smaller than females in all measurements (see paper for details).
  • Measurements suggest that the Chiloé population includes rufa Red Knot.
  • Weights of birds were higher at the end of April than at the start of March, suggesting an increase in body mass of between 2.9 and 3.6 grammes per day; figures that are comparable to other studies of Knot.
  • In spring, marked birds were reported in Peru, the Gulf of Mexico, Minnesota and Manitoba. These last two sightings are on the Mid-Continental Flyway, which is used by waders heading for both Alaska and Northern Canada, as might be expected of roselaari and rufa Red Knot, respectively.
  • The red-flagged birds pictured shown here were photographed by Peter Bergeson (above right) and Jean Hall (below) in South Carolina and Florida, respectively) clearly suggesting that they are rufa Red Knot.

Conservation implications

Chiloé is an important non-breeding area, where Red Knot fuel up for non-stop 8,000 km flights to the Gulf of Mexico, one of the longest migration legs for the species. Now that Gherardi-Fuentes et al have shown that these flocks include ‘Endangered’ rufa, it makes sense to provide some designated protection to the Chiloé Archipelago population of Red Knot. You can download the current Conservation Plan for Migratory Shorebirds in Chiloé.

This relatively small-scale study of Red Knot has emphasised two important points about shorebird conservation.

  • The protection of sites that hold important populations of key species provides benefits for other waders that use similar habitats. In this case, sites designated for Hudsonian Godwits and Whimbrel are being used by two subspecies of Red Knot, at least one of which is ‘threatened’.
  • Waders from one breeding population use a range of sites when migrating and during the ‘wintering’ period. Given that it is hard to know all of the possible sites that link to one breeding area, it is pragmatic to protect as many different sites as possible, across a broad range of countries. There is more about this in Spoon-billed Sandpiper: Track & Trace.
Caulín Bay in Chiloé

Paper

It is interesting that we are still discovering important information about the origins of population of Red Knot, a species that has been at the heart of shorebird research for decades. Will genetic techniques and tracking reveal more surprises? And, more intriguing, how much more is still to be discovered about less well-studied species?

Here’s a link to the paper in Bird Conservation International:

Insights into migratory connectivity and conservation concerns of Red Knots Calidris canutus in the austral Pacific coast of the Americas. Camila Gherardi-Fuentes, Jorge Ruiz and Juan G Navedo (2021).

Further work on the genetics of American Red Knot

Research published later in 2021 reinforces the ‘every wintering site is important’ message in the paper by Camila Gherardi-Fuentes and colleagues.

As indicated earlier, the rufa subspecies of Red Knot is listed as “Threatened” or “Endangered” in the USA, Argentina and other countries in the flyway. Recent research by Yvonne Verkuil, Patricia González and colleagues has involved genotyping 150 Red Knot, to test whether rufa birds spending the boreal winter in Argentina (Tierra del Fuego and Río Negro), northern Brazil (Maranhão) and south-eastern USA (Florida) can be considered to come from one interbreeding population. The team detected genetic differences that they argue warrant the recognition of three nonbreeding regions in Argentina, northern Brazil and south-eastern USA (each hosting 3,600–13,000 Red Knots) as distinct units, even though the breeding origins of the birds remain unknown. The unique assortments of genotypes suggest that these regions receive birds from different areas within the Arctic, meaning that each site needs to be considered as an important (and irreplaceable) part of the Red Knot puzzle. Details of the paper:

Genetic structure in the nonbreeding range of rufa Red Knots suggests distinct Arctic breeding populations. Yvonne I Verkuil, Erika Tavares, Patricia M González, Kristen Choffe, Oliver Haddrath, Mark Peck, Lawrence J Niles, Allan J Baker, Theunis Piersma & Jesse R Conklin. Ornithological Applications. Volume 124, 2021, pp. 1–11. https://doi.org/10.1093/ornithapp/duab053


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.

Oystercatcher Migration: the Dad Effect

What determines whether some birds migrate and others do not? This question is fundamental to understanding how migratory systems change over time but the causes of individual migratory behaviours have proved difficult to isolate.

Verónica Méndez and colleagues are studying Icelandic Oystercatchers, some of which remain in Iceland for the winter but most of which migrate across the Atlantic to Ireland, Britain and mainland Europe. In a 2021 paper in Scientific Reports they show that a chick’s migratory behaviour seems to align with the behaviour of its father but not its mother. What can explain this pattern?

The story so far

The Icelandic Oystercatcher study system has already featured in three WaderTales blogs. The first was Migratory decisions for Icelandic Oystercatchers. This focused upon the key questions that Verónica Méndez and colleagues from the universities of Iceland, East Anglia (UK) and Aveiro (Portugal) are trying to answer.

  • Why do some Oystercatchers migrate when others don’t?
  • Is it the same birds each year?
  • Do resident or migrant birds have an advantage when it comes to choosing a territory and raising chicks?
  • Do chicks follow the same migratory patterns as their parents?

When the first blog was written, in 2015, eight colour-ringed Oystercatchers had been seen in Ireland and the UK, and five had been seen wintering in Iceland. Fast forward to the next blog in 2018 – Mission impossible? Counting Iceland’s wintering Oystercatchers – where counts showed that over 11,000 Oystercatchers spend the winter in Iceland. Using colour-ring sightings of resident and migratory birds, the research team concluded that this total is about 30% of the whole Icelandic population. The other 70% fly south across the Atlantic each autumn, with no individuals yet observed to change what they do between years.

In the third blog – Which Icelandic Oystercatchers cross the Atlantic? – some patterns were starting to emerge.

  • Females and males are equally likely to migrate.
  • Size does not matter – small and big birds are equally likely to migrate
  • There are regional patterns across Iceland, with birds breeding in the west being most likely to be resident.
  • Birds do not pair up assortatively – residents don’t pair up with other residents before the migrants return, for instance.

Family ties

In most species of waders, parents protect their chicks and take them to suitable feeding areas but they do not actively feed them. Parental care in European Oystercatcher includes foraging for food and bringing it back to the chicks. This is why it is possible for Oystercatchers to nest on the roofs of buildings (Oystercatchers: from shingle beach to roof-top), where they are out of the reach of ground predators.

Focusing on chicks

To be able to understand the relationship between migratory behaviour in adults and their chicks, you need to be able to mark and then attempt to follow all of the members of a family. Adult Oystercatchers generally keep the same mates and nest in the same areas year after year, enabling the establishment of marked population of birds in different parts of Iceland. Between 2015 and 2018, a total of 615 incubating adults were caught. By following the outcomes of nesting attempts and then monitoring the growth of chicks, the research team also managed to individually mark 377 chicks.

Three colour-ringed chicks. Where will they go?

The success of the whole project relied heavily upon winter sightings of marked birds within Iceland and in Ireland, the UK and continental Europe. Through a network of volunteer observers reporting sightings of marked individuals across the wintering range, the migratory behaviours of 227 of the 615 colour-marked adults and 50 of the 377 colour-marked chicks had been identified at the time that this paper was written. In addition, it was possible to infer the migratory behaviour of 353 marked adults using measurement of isotope ratios (δ13C and δ15N) of feathers that were grown in the winter (as described here).

The analyses in the paper by Verónica Méndez and her colleagues are based upon 42 marked chicks of parents for which the migratory behaviour of both parents is either known or can be inferred from isotopic signatures. These chicks all fledged successfully and were seen during the winter period, either in Iceland or having crossed the Atlantic. In three cases, two chicks from the same broods are known to have behaved in the same way. More data have become available since the analyses, all confirming the same patterns.

Results

It is possible to imagine a scenario in which late or slow-growing Oystercatcher chicks might be more likely to stay in Iceland than their more mature counterparts – simply by developing too late to gain enough resources to cross the Atlantic. Analysis of hatch dates and growth parameters did not suggest the existence of such a link, as described in the paper.

This young Oystercatcher was spending its first winter on the coast of western Iceland

The interesting finding of this study is the link between the behaviour of parents and chicks. Data generated by observations of colour ringed individuals (adult and chicks) and from isotopes (adults) established 21 chick/parent associations.

  • Of the sixteen chicks raised by migrant mothers, eight migrated and eight remained in Iceland.
  • Of the five chicks raised by resident mothers, three migrated and two remained in Iceland.
  • All ten of the chicks raised by migrant fathers migrated from Iceland.
  • Of the eleven chicks raised by resident fathers, one migrated and ten remained in Iceland.
  • Seven chicks that fledged from pairs with one resident and one migrant parent adopted the migratory behaviour of the father.

This is pretty compelling evidence that chick migratory behaviour is associated with paternal (and not maternal) migratory behaviour!

What does this mean?

There is no evidence of genetic control of migratory destinations and both Oystercatcher parents care for chicks, so what mechanism could produce such strong paternal but not maternal effects?

The authors suggest that the migratory behaviour of individual oystercatchers may be linked to social interactions they experience during the post-fledging period. In shorebird species, such as Oystercatchers, mothers commonly depart before the chicks fledge, or at about the same time. Fathers often provide parental care for longer and this extended period of the parental bond may underlie the link between paternal and juvenile migratory behaviour in Icelandic Oystercatchers. Despite being able to fly and feed independently, juvenile Oystercatchers in Iceland have been seen begging for food several months after fledging, suggesting that some parents (most likely fathers) may care for youngsters much longer than in other species.

This Iceland-ringed Oysterctatcher was photographed in Guernsey in January 2021. It departs at the start of February each year.

Under this extended-care system, a chick that is being look after by a resident male may well become a resident, simply by following dad. As autumn arrives, the youngster can follow his parent when he moves to the coastal mudflats where resident Icelandic Oystercatchers spend the non-breeding season. Autumn turns to winter and the chick is destined to be a resident.

Is it possible to explain a similar link for migrants? As the breeding season comes to an end, migrant fathers leave their breeding areas and head south, across the Atlantic, leaving fledged youngsters to fend for themselves. Groups of youngsters gather together in flocks which also include adults that are feeding up in preparation for migration. Although not influenced by their own fathers, chicks may follow the cues of other migratory adults, thereby creating the patterns seen in this paper.

Most of the chicks included in these analyses were early-fledged birds, simply because earlier nesting attempts tend to be more successful. The research team were unable to detect any significant effect of fledging date on migratory behaviour but they do not rule out the possibility that late-fledging individuals lack the time or resources to undertake a migratory journey, irrespective of paternal behaviour.

The broader context

Migratory behaviour typically arises in seasonal environments, allowing individuals to exploit peaks of resource abundance in distinct locations across the world. Rapid shifts in the distribution and migration phenology of many migratory species present challenges to site-based conservation strategies. There is an urgent need to understand the processes that influence individual migratory behaviour, in order to attempt to predict species’ responses to environmental change.

The findings in this paper suggest that the social interactions experienced by individuals can directly influence the development of their migratory behaviour, and that the extent and timing of parental care may be key in shaping individual access to these social interactions. You can read the full paper here:

Paternal effects in the initiation of migratory behaviour in birds Méndez V., Gill, J.A., Þórisson, B., Vignisson, S.R., Gunnarsson, T.G. & Alves J.A.


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.

Winter conditions for Whimbrel

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.

Crabs provide a large part of a Whimbrel’s winter diet

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.

A colour-ringed bird hiding in a Bijagós flock

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.

Trying to find Whimbrel in a sandstorm in Banc d’Arguin

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.

Life in and amongst the mangroves of the Bijagós

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.

Carry-over Effects

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.

Paper

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

This Whimbrel, photographed in Bijagós is wearing colour-rings that were fitted in Iceland

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.

Further reading

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.