January to June 2022

Here are brief summaries of the first nine WaderTales blogs of 2022. As ever, I am grateful to the authors of the papers that underpin the blogs; they have worked with me to make sure that I get the stories right! I have not covered every new paper; perhaps your favourite is in the pipeline or perhaps I did not happen to spot it? The blogs are described in order of publication.

Welsh Oystercatchers

To explain how flexible Oystercatchers can be in response to changes in their food supply, Katharine Bowgen has brought together long-term data collected by wader ringers and WeBS counters, and added in annual assessments of cockle stocks on the Burry Inlet (South Wales). This paper has a particular resonance, as I remember teaching students about the Burry Inlet Oystercatcher controversy of the 1970s, when complaints from shellfishers led to the deaths of thousands of birds. We understand more about the relationship between shellfish stocks and bird numbers now but what happens when Oystercatchers can’t find food? This paper makes a strong case for the protection of networks of sites, so that individuals have alternatives when needed.

Australian stock-take

One of the great joys of writing WaderTales blogs is that I get to ‘visit’ the flyways of the world without having to burn carbon. How many shorebirds use the East Asian-Australasian Flyway? is a flyway-wide stock-take of the waders that visit Australia and New Zealand, led by Birgita Hansen. It is shocking that a flock of 350 Far-eastern Curlew now constitutes 1% of the global population and that the population of Curlew Sandpipers has halved in double-quick time, but the key strength of the paper is the clear explanation of a methodology that can be used in the future, to monitor changes in numbers.

Chick vocalisation

Big analyses of data sets are very important but it’s lovely when you learn more about the natural history of species that birdwatchers know well. In Australia, Kristal Kostoglou recorded the calls of the chicks of Red-capped Plovers and Southern Masked Lapwings, that were being ringed and measured in the hand. In Chick squeaks I describe how calls get deeper with age, which is not surprising, but that the calls of males and females can become distinguishable from a very early age. Male Red-capped Plover chicks are more demanding than their sisters!

Trans-oceanic migration

There have been several recent wader papers that interpret data obtained from birds when on migration. One of the interesting questions being asked is, “Do shorebirds account for wind displacement continuously or correct for drift later?”. Navigating a vast ocean summarises Jenny Linscott’s work on Hudsonian Godwits, as they cross the Pacific Ocean and then the Gulf of Mexico, on their way from Chile to Alaska. She and her fellow authors show that flocks make continuous adjustments, demonstrating that birds ‘know where they are’ and giving them the ability to fly extremely long distances without running out of energy. There’s some clever maths too!

Hiding in the trees

In the second paper from her PhD, Triin Kaasiku looks at the breeding success of Estonian coastal waders that nest at different distances from woodland. Keep away from the trees describes these ‘edge effects’. In a part of the world where waders are in diminishingly short supply, hatching success is six time as high in open areas as in areas that are within one kilometre of forest edge. The Baltic coast used to be a haven for species such as Curlew and Dunlin but reduced grazing and forestry plantations have provided hiding places for predators. Alongside increased predation, breeding waders are also having to contend with an increasing numbers of nest inundations, arising from summer storms.

Curlew hunting

Curlew hunting stopped in Great Britain in 1982, when the declining wintering population received protection under the new Wildlife & Countryside Act. A fascinating paper by Ian Woodward and BTO colleagues teases apart the positive effects of the cessation of shooting and more benign winter weather. It is summarised as Curlew: after the hunting stopped.

I am old enough to remember when Curlew were hunted in East Anglia. The pâté made from autumn-shot birds is reputed to have been very tasty; I recall Clive Minton getting back in his land-rover and reporting that he had been offered some, when asking for permission to cannon-net Curlew on a Norfolk land-owner’s estate.

Personal appreciation of Whimbrel

On 27 April, Jenny Gill and I were at Eyrarbakki, on the south coast of Iceland. As we watched, small groups of Whimbrel were coming in off the sea. Others were resting on the seaweed-covered rocks, a few were feeding and some flew straight by. Watching waders arrive in Iceland is always magical but, from sightings of satellite-tagged Whimbrel, we could be pretty sure that these tired birds had just completed five-day, direct flights from west Africa. I could not wait to get back to base and to share our observations. It was a good excuse to round up the Whimbrel stories in other WaderTales blogs, as you can read in Whimbrels arrive in Iceland.

Power-line problems

We have seen huge changes in Iceland, since we first visited in 2000, but how are these affecting shorebirds? In the first paper of her PhD (Effects of land conversion in sub-arctic landscapes on densities of ground-nesting birds), Aldís Pálsdóttir investigated how distributions of breeding waders are affected by power-lines. She discovered significantly depressed numbers several hundred metres from the transmission lines, with Whimbrel and Redshank being the most obviously impacted. Her results are written up as Power-lines and breeding waders. With an increasing global reliance on electricity, these are important findings for planners and conservationists.

Conflict with forestry

Just a few days later, Aldís Pálsdóttir’s second paper was published. By mapping distributions of breeding waders in the vicinity of forests, she has shown that new plantations have a massive effect on distributions. In lowland Iceland, the most vulnerable species appear to be Dunlin and Oystercatcher, followed by Whimbrel, Black-tailed Godwit and Golden Plover. It should be noted that three-questers of Europe’s Whimbrel nest in Iceland, as well as half of the Golden Plover and Dunlin. Aldís and her fellow authors argue that Iceland’s waders need a strategic forestry plan. They estimate that recently-planted woodland and forests have already removed the breeding territories of tens of thousands of waders.


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.

Iceland’s waders need a strategic forestry plan

More and more trees are being planted in lowland Iceland – and further increases are planned, in part encouraged by the suggestion that this will mitigate for climate change. Forestry is potentially bad news for Whimbrel, Black-tailed Godwit and other waders that breed in open habitats, and which migrate south to Europe and Africa each autumn. Are there ways to accommodate trees while reducing the damage to internationally important populations of waders?

Pressure on Iceland’s breeding waders

Iceland is changing; more people want second homes in the countryside, the road network is being developed to cope with more and more tourists, new infrastructure is needed to distribute electricity, agriculture is becoming more intensive and there is a push to plant lots more trees. The south of the country is seeing the most rapid loss of open spaces, providing opportunities to study how these incursions affect ground-nesting species, particularly breeding waders.

One of the big changes, especially in Southern Iceland, has been the planting of non-native trees, as shelter belts around fields and country cottages and, more significantly, as commercial crops. Iceland has been largely treeless for hundreds of years but climatic amelioration has facilitated rapid forestry development in areas where tree growth was previously limited by harsher environmental conditions. Seeds of some non-native species are blown on the wind for a kilometre or more, to germinate in open land, well beyond the edge of planned forests.

Most of the new forests are in lowland areas, where we also find the most important habitats for many ground-nesting bird populations. Lodgepole pines may be good news for Goldcrest and Crossbills but not for species such as Golden Plover, Dunlin & Redshank. For breeding waders, the most obvious impact of a new forest is direct loss of breeding habitat but trees can have wider effects, by providing cover for predators and breaking up swathes of open land that are used at different stages of the breeding season. Little is currently known about how predators in Iceland use forest plantations but any perceived risks of predator presence and reduced visibility is likely to influence densities of birds in the surrounding area.

Iceland’s open habitats have suited breeding waders for hundreds of years

Aldís E. Pálsdóttir’s studied changing bird populations in lowland Iceland during her PhD at the University of Iceland, in collaboration with researchers from the University of East Anglia (UK) and the University of Aveiro (Portugal). Among the most concerning of these changes is the rapid expansion of forestry in these open landscapes.

Assessing the potential impacts of trees

In a 2022 paper in the Journal of Applied Ecology, Aldís assesses whether densities of ground-nesting birds are lower in the landscape surrounding plantations and whether these effects vary among plantations with differing characteristics. She and her fellow authors then quantified the potential impact of differing future afforestation scenarios on waders nesting in lowland Iceland.

Forestry currently covers about 2% of Iceland’s land area so the potential for growth is massive. In 2018, the Icelandic government provided additional funding to the Icelandic forest service to increase the number of trees planted, with a goal of enhancing carbon sequestration. As forestry primarily operates through government grants to private landowners, who plant trees within their own land holdings, plantations typically occur as numerous relatively small patches in otherwise open landscapes. These features make Iceland an ideal location in which to quantify the way that plantations affect densities of birds in the surrounding habitats, and to identify afforestation strategies that might reduce impacts on globally important wader populations.

To measure the effects of plantation forests on the abundance and distribution of ground-nesting birds, in particular waders, 161 transect surveys were conducted between May and June 2017. To avoid systematic bias arising from possible “push effects” of corralling birds in front of the surveyor, surveys were conducted along transects that started either at the edge of the plantation, with the observer moving away (79 transects), or started away from the plantation, with the observer walking towards it (82 transects). Please see the paper for the full methodology. The variation in density with distance from plantation was used to estimate the likely changes in bird numbers, resulting from future afforestation plans, and to explore the potential effects of different planting scenarios.

Bird communities change around plantations

Snipe densities are highest close to young forests

On the transects, 3713 individual birds of 30 species were recorded. The nine most common species (excluding gulls, which rarely breed in the focal habitats) were seven waders (Oystercatcher, Golden Plover, Dunlin, Common Snipe, Whimbrel, Black-tailed Godwit & Redshank) and two passerines (Meadow Pipit & Redwing). These species accounted for 88% of all birds recorded.

  • Of the seven waders, Snipe was the only one found in significantly higher numbers closer to plantations. Snipe density declined by approximately 50% between the first (0-50 m) and second (50-100 m) distance intervals, suggesting a highly localised positive effect of plantations on Snipe densities.
  • Densities of Golden Plover, Whimbrel, Oystercatcher, Dunlin and Black-tailed Godwit all increased significantly with increasing distance from plantations. Dunlin and Oystercatcher showed the largest effect (~15% increase per 50 m), followed by Whimbrel (~12%), Black-tailed Godwit (~7%) and Golden plover (~4%).
  • Although Redshank did not show a linear relationship with distance from plantation edges, densities were lowest close to the plantation edge.
  • There were more Redwings close to woodland edges but Meadow Pipit showed no change in density with distance from plantations.

Golden Plover, Whimbrel and Snipe were found in lower densities close to the tallest plantations (over 10 m), when compared to younger plantations (tree height 2m to 5m), suggesting that the impact of forests gets more pronounced as the trees grow. Plantation density and diameter had no additional effect on the species that were in lower densities closer to the plantations, implying that the mere presence of plantations induces the observed changes in abundance. See the paper for more details.

The bigger picture

Aldís Pálsdóttir and Harry Ewing walked every step of every transect and made detailed counts of what they saw – data that are invaluable when considering local impacts of plantations – but the paper becomes even more interesting when the authors look at the bigger picture. When plantations are distributed across these open landscapes, in different configurations, what will be the accumulated effects on the numbers of breeding waders? They estimate likely changes in abundance resulting from planting 1000 ha of plantation in different planting scenarios, ranging from a single block to lots of small patches.

  • Planting 50 smaller patches of 20 ha, instead of 1000 ha of forest in one large patch, is estimated to double the resulting decline in abundance (because there is more forest edge and hence a bigger effect on more open habitat)
  • This effect increases even further as the patches become smaller; in their models, planting 1000 blocks each of 1 ha would have nine times the impact of planting one forest of 1000 ha.
  • Proximity of woodland seems to be the driver of local distributions of breeding waders so the authors suggest that the amount of edge (relative to area) should be minimised, to reduce the impact of a plantation – which means making forests as near circular as possible.

It is clear that fewer larger forestry plots are likely to be less bad than lots of small, local plantations, in terms of the effects on wader populations. The figure below illustrates how much more land is affected when one woodland is replaced by four with the same total area. The grey area (equivalent to a 200 metre annulus) accounts for 88 hectares in the one-patch illustration and 113 hectares for four patches.

An urgent need for action (and inaction!)

Iceland holds large proportions of the global nesting populations of Golden Plover (52%), Whimbrel (40%), Redshank (19%), Dunlin (16%) and Black-tailed godwit (10%) (see Gunnarsson et al 2006) and is home to half or more of Europe’s Dunlin, Golden Plover and Whimbrel. Data in the table alongside have been extracted from Annex 4 of the report, which was discussed at the 12th Standing Committee of AEWA (Agreement on the Conservation of African-Eurasian Migratory Waterbirds) in Jan/Feb 2017.

Aldís measured the areas of 76 plantations in her study, using aerial photographs. The total area of woodland was about 2,800 ha and the total amount of semi-natural habitat in the surrounding 200 m was about 3,600 ha. Using the reduced densities that she found on the transects and the direct losses for the plantations themselves, she estimates potential losses of about 3000 breeding waders, just around these 76 forest plots. Extrapolating this figure to the whole of the Southern Lowlands of Iceland, the total losses resulting from all current plantations are likely to already be in the tens of thousands. Worryingly, the densities measured on the transects in this paper (even 700 m from forest edge) were well below those measured (slightly differently) in previous studies of completely open habitat, suggesting that losses may already be significantly higher than estimated in the paper.

A scary statistic in the paper is that “6.3% of the Icelandic lowlands is currently less than 200 m from forest plantations”. Given the incentives to plant lots more trees, this is particularly worrying for species such as Black-tailed Godwits, the vast majority of which breed in these lowland areas (between sea level and 300 metres).

Non-native trees are spreading beyond the boundary of a planned forest

It has been suggested that breeding waders might move elsewhere when impacted by forestry but migratory wader species are typically highly faithful to breeding sites. If birds are not going to move to accommodate trees, then perhaps plantations should be located where bird numbers are naturally low, such as in sparsely or non-vegetated areas, at higher altitudes and on slopes? Planning decisions could usefully be informed by surveys of breeding birds, to identify high-density areas that should be avoided.

The severe impact that planting forests in open landscapes can have on populations of ground-nesting birds emphasises the need for strategic planning of tree-planting schemes. Given Iceland’s statutory commitments to species protection, as a signatory to AEWA and the Bern Convention on the Conservation of European Wildlife and Natural Habitats, and the huge contribution of Iceland to global migratory bird flyways, these are challenges that must be addressed quickly, before we see population-level impacts throughout the European and West African Flyway.

To learn more

The take-home message from this work is clear. Local planning decisions and the ways in which forestry grants are allocated are producing a patchy distribution of plantations across the lowlands of Iceland, and this is bad news for breeding waders.

The paper at the heart of this blog is:

Subarctic afforestation: effects of forest plantations on ground-nesting birds in lowland Iceland. Aldís E. Pálsdóttir ,Jennifer A. Gill, José A. Alves, Snæbjörn Pálsson, Verónica Méndez, Harry Ewing & Tómas G. Gunnarsson. Journal of Applied Ecology.

Other WaderTales blogs that may be of interest:

Forest edges

Work by Aldís Pálsdóttir (pictured right)

Changing agricultural systems in Iceland (work by Lilja Jóhannesdóttir)


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.

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.

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.

A Rhapsody of Whimbrel

Around the globe, the seven-note whistle of the Whimbrel is a spring theme tune for shorebird migration. Where are flocks going to, as they gain height and head north in March, April and May, and why have birds chosen that particular moment to depart?

In a paper in Frontiers in Ecology & Evolution, Camilo Carneiro, Tómas Gunnarsson and José Alves analyse individual migration tracks of birds flying from West Africa to Iceland, including journeys made in successive springs by birds that have been part of the same study for several years. Can these birds help to explain how Whimbrel use time and weather cues in their travel ‘planning’ and might their plans change during the course of their lives?

Look up to the skies and see

Almost all of the Whimbrel that nest in Iceland migrate to West Africa after the breeding season, with just a tiny number wintering in Europe.

Life has just begun

The Whimbrel’s story starts in Southern Iceland. It would be great to tag chicks when they hatch, in order to learn what happens to juveniles in their first autumn, as they head south without parental guidance. For the moment, however, Camilo Carneiro, José Alves and Tómas Gunnarsson are focusing on parent birds which they have been catching on their nests since 2012.

The research summarised in the paper at the heart of this blog uses data from 66 retrieved geolocators that have been carried by a total of 39 individuals.  Light data, temperature readings, conductivity and the timing of contacts with water all help to determine the start and end points of individual flights.

Any way the wind blows?

As the authors write in the summary of the paper, “Weather conditions are important during migration, particularly wind and temperature, and can play a crucial role in the timing of events during the annual cycle of migratory birds.” Camilo and his colleagues have investigated how wind conditions, temperature and spring departure date may drive individuals to pursue either a ‘stop-over’ or ‘direct flight’ strategy in the spring. They suggest that one or two of the following scenarios might be envisaged:

Competition on the breeding grounds can be quite vigorous
  • Whimbrels may make migratory decisions prior to departure, given local weather conditions. If migratory behaviour is defined prior to departure then that would suggest that departure time from West Africa may not be associated with the wind conditions that can assist individuals on the first stage of their journeys.
  • Whimbrels may adjust migratory behaviour during flight, depending on conditions experienced en route. As birds fly north, from 37°N (level with the Mediterranean) to 50°N (SW tip of England), they can make decisions to stop off, rather than continue straight to Iceland. Potentially, they could assess the amount of support they have been receiving from favourable winds and assess the positive effect on fuel reserves. Temperature could also act a cue to weather conditions in Iceland.

Little high, little low

Wader biologists and birdwatchers will have seen flocks of waders preparing for departure from breeding, stop-over and wintering locations.  Birds become restless and groups of birds start to circle and rise into the sky.  The first, departing flock may be joined by others, taking off in small groups to catch up with birds that seem to have a plan. At the same time, birds can also be seen dropping out of the flock to wait for another day.

Once formed, a migratory flock will be made up of individuals that may be heading for points that are hundreds or (for some species) even thousands of kilometres apart but will fly together for this next leg of the journey.

Tómas Gunnarsson has seen Whimbrel circling and gaining height at departure from the south coast of Iceland in autumn, suggesting that they may assess wind conditions at different altitudes before making a final decision and setting their course south. It seems plausible that the same sort of thing might happen during spring migration from West Africa. At some stage, with a large number of tracked birds, perhaps it will be possible to study if waders sample wind speed and direction prior to departure and whether decisions made at this point affect the amount of support they receive from wind conditions when on migration. There’s a great paper about how wind affects the migration of much larger Honey Buzzards by Wouter Vansteelant.

Once in the air, birds in a migrating flock can be impeded by headwinds, assisted by tailwinds and pushed laterally by crosswinds. As has been shown when tracking trans-Pacific journeys of Bar-tailed Godwits, storms and pressure systems can provide a sling-shot effect to accelerate progress, cause birds to temporarily abort their journeys or even back-track a thousand kilometres, in order to refuel for another migration attempt.

Nothing really matters

Camilo and his colleagues compared 9 direct flights with 48 that included stops in Portugal, France, Ireland and north-west Britain. Birds that departed later tended to undertake direct flights. Interestingly, they found little evidence that conditions were linked to an individual’s migration option:

  • For individuals undertaking either direct flights or stopovers, the wind conditions at departure did not differ from winds during the previous seven days, suggesting little or no selection for wind conditions to initiate the flight.
  • Individuals on direct flights encountered similar winds as they passed through the region 37°N to 50°N (Mediterranean to SW tip of England) as those on stop-over journeys.

Too late, my time has come?

For late-departing birds that fly straight to Iceland, there is the potential to catch up with (and even overtake) birds that left earlier and subsequently stopped in countries such as Ireland. In such circumstances, one could imagine that birds still in West Africa in late spring may ‘opt’ for a direct flight. However, the wind patterns over a period of seven days prior to departure were no different to the ones at the start of a non-stop flight. There did not seem to be any weather-related reason for these birds delaying departure as long as they did.

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Camilo Carneiro is also investigating the breeding success associated with different arrival times and migration patterns

For birds that do stop-off en route in Britain and Ireland, which are relatively close to the breeding sites in Iceland, there are potential benefits, as discussed in more detail in the paper:

  • Individuals might be able to assess (remotely) the weather conditions at their breeding sites if there are links between weather patterns in Iceland and those in Ireland and western Britain. If this is the case, then they can adjust the start of the final Atlantic crossing so as to arrive in Iceland when conditions are likely to be favourable.
  • Whimbrels can feed up in Britain & Ireland, potentially arriving in Iceland in better condition than birds that have flown directly from West Africa.

Carry on, carry on

The fascinating thing about being able to track individuals is that we are starting to understand the conditions that may lead to the migration patterns we see at the population level. In other WaderTales blogs, some of the key processes and patterns have already been discussed:

Given this body of knowledge, suggesting little flexibility once an individual’s phenology is fixed, one might expect that an individual Whimbrel, travelling from West Africa to Iceland, would adopt the same migration pattern in subsequent years. Tagged Whimbrels, for which there were repeat migration tracks, mostly ‘did the same’ but not always.

Any way the wind blows, doesn’t really matter

The key finding of Camilo Carneiro and his colleagues is that weather conditions are not the main driver of different spring migratory timings in Icelandic whimbrels. As suggested in Whimbrel: time to leave (and the paper Why are Whimbrel not advancing their arrival dates into Iceland?) departure from West Africa is the most firmly scheduled point in a Whimbrel’s annual cycle. However, three birds provide an opportunity to build a modified hypothesis.

In the course of this study of Icelandic Whimbrel, thirteen birds have provided multiple years of data and there have been three cases in which birds have switched migratory behaviour, seemingly contradicting the ‘birds always do the same thing’ theory. In each case, individuals changed from ‘direct-flight’ to ‘stop-over’ and, having changed, they then repeated the ‘stop-over’ option. The fact that all three individuals made the same change hints towards individual refinement of behaviour. Since there appears to be no clear advantage to direct migration, perhaps individuals are moving to a two-stage migration when they discover that this is a possibility?

Is this just fantasy?

This is where the paper stops but it will be fascinating if the research can continue:

  • Are changes always going to be in the same direction – from ‘direct-flight’ to ‘stop-over’?
  • Most juveniles are thought to fly directly to West Africa. By tracking young birds, caught in Iceland prior to departure, perhaps it will be possible to work out the proportion of first journeys that are direct?
  • Young birds that fly south across the Atlantic will not have learnt that there is any land between Iceland and West Africa. Do they retrace their journeys when they fly north again, eighteen months later?
  • What proportion of these birds that start on the ‘direct-flight’ strategy later switch to ‘stop-over’?
  • If some juveniles stop off in western Europe, on their first journeys south, perhaps these birds also stop off on their first journeys north? That might mean that they never used the ‘direct-flight’ option.
  • And how do flocks work? Will it ever be possible to deploy enough trackers to see whether ‘direct-flight’ individuals learn that there’s an alternative option by travelling in flocks with birds that use the ‘stop-over’ strategy.

It’s all fascinating stuff about the Queen of waders!

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Anyone can see

The paper featured in this blog is: Linking weather and phenology to stopover dynamics of a long-distance migrant by Camilo Carneiro, Tómas G. Gunnarsson & José A. Alves, published in Frontiers in Ecology & Evolution.


GFA in Iceland

WaderTales blogs are written by Graham Appleton, to celebrate waders and wader research.  Many of the articles are based on previously published papers, with the aim of making wader science available to a broader audience.

@grahamfappleton

Where to nest?

pic whimbrelThere is nothing more obvious than an Oystercatcher sitting on his or her nest, but a brooding Snipe can be invisible until almost trodden upon. Which strategy works better: nesting in plain view but laying cryptically camouflaged eggs or hiding yourself and your nest in a clump of grass? Which species is most likely to hatch a successful brood of chicks and in what circumstances? In a 2020 paper in IBIS, Becky Laidlaw and colleagues analysed nest site characteristics and nest locations of 469 wader nests in Iceland in order to provide some answers

The perils of ground-nesting

pic hatching whimbrel

Hatching Whimbrel eggs, with the tell-tale shell fragments that signal a nesting attempt has been successful

Almost all waders are ground-nesters, which makes them highly vulnerable to a wide range of nest predators. To reduce the risks of predation, different strategies have evolved. In some species, nests are placed out in the open, and the camouflage is provided only by mottled egg colouration that resembles the background. In other species, nests are secreted in vegetation, meaning eggs and incubating adults are concealed from predators.

In both groups of species, the risk of nests being predated might vary, depending on the surrounding habitat. For open-nesting species, for example, clutches that are laid in large patches of similar habitat may be harder for predators to locate. The same could apply to closed-nest species that hide their nests; Snipe nests may be tricky to find in extensive areas of long grass but perhaps more at risk if there are only a few suitable clumps of long grass that predators need to check out.

pic hidden Redshank

Iceland: a wader factory

tableAs discussed in previous WaderTales blogs, particularly Do Iceland’s farmers care about wader conservation? Iceland is hugely important as a European ‘wader factory’. As farmland elsewhere has become less suitable for species such as Redshank and Snipe, the global importance of the country has increased (see table alongside for most recent figures from an AEWA report)  With this in mind, it is important to understand the factors that underpin the population dynamics of Iceland’s breeding waders.

Working in South Iceland, Becky Laidlaw and her co-authors tried to find as many nests as possible during the summers of 2015 and 2016. This area is largely a mosaic of open habitats, although there are more patches of forestry than there were twenty years ago. Most of the Southern Lowlands area is farmed, on a gradient between intensive and semi-natural, and this is reflected in the distribution of breeding waders (see Farming for waders in Iceland).

pic rope

Dragging a light rope across the vegetation to flush nesting birds

For this project, nests were located by surveys from vehicles and on foot, through observation of incubating adults, systematic searching, incidental flushing of incubating adults and rope-dragging (dragging a 25 m rope, held between two fieldworkers, lightly across vegetation) to flush incubating adults.

The analysis in the resulting paper in IBIS focuses on 469 nests of three open-nesting species (Oystercatcher, Golden Plover and Whimbrel) and three species that hide their nests in tall vegetation (Redshank, Snipe and Black-tailed Godwit). The team recorded the habitat and vegetation structure around each nest (at the nest, within a 5 m x 5 m square and in a wider 50 m x 50 m square) and worked out which nests hatched successfully and which were predated. The date and time of predation were determined, where possible, with nest-cameras providing extra information for some nests. Cameras captured nest-predation events involving Arctic foxes, Arctic Skuas, Ravens and sheep.

Interestingly, 2015 and 2016 were very different wader breeding seasons. The graphic below shows the mean temperatures for the months from April through to July (encompassing the wader breeding season at this latitude) were much cooler in 2015 than in 2016, representing average monthly difference of between 1.5°C and 2.5°C. At high latitudes these figures translate into very different rates of vegetation growth.

pic pretty graph

First, find your nest

When nests were first located, their positions were marked and referenced using GPS. Eggs were floated in water to provide an estimate of laying date and thereby predict hatching date. As the chick develops within an egg, the density of the egg falls. A newly laid egg will lie on the bottom of the flotation vessel. Over the next few days the ‘blunt end’ rises until the egg is still touching the bottom but vertical. Eggs in the late-development stage float ‘point-end-down’, with the latest eggs floating at an angle to the vertical (method described by Liebezeit et al.).

pic skua-ed goldie eggs

This Golden Plover nest was probably predated by an Arctic Skua

Nests were considered successful if one or more eggs hatched, and predated nests were defined as those that were empty in advance of the predicted hatch date or those without any eggshell fragments in the nest (a sign of successful hatching). To determine the time and date of nest failures, iButton dataloggers were placed in a randomly selected subsample of nests. These loggers recorded a temperature trace every ten minutes. A sharp and permanent decline in nest temperature below incubation temperature indicates nest predation. In both study years, motion-triggered cameras were deployed on a sample of open-nesting species to determine the predator species active on these nests.

When each nest was first located, the percentage of eggs visible from directly above the nest was estimated and the habitat surrounding each nest was assessed in the field at three spatial scales: the nest cup, the 5 m x 5 m and the 50 m x 50 m area surrounding each nest. Details are in the paper.

Which nests survive through to hatching?

Over the breeding seasons of 2015 and 2016, the outcomes of 469 wader nests were assessed. 259 hatched successfully (55%), 192 were predated (41%), 13 were abandoned, 7 were trampled and 2 were mown. A nest-loss rate of 40% is fairly typical for ground-nesting waders, when compared to studies in different countries and habitats.

pic fox attack

Daily nest predation rates did not vary significantly in relation to the habitat heterogeneity or the extent to which the dominant habitat covered the area surrounding the nest, at either 5 m x 5 m or 50 m x 50 m scales. Most clutches were laid in habitats that were the same or similar to the surrounding areas. Where there were differences, the dissimilarity between the habitat at the nest cup and in the surrounding area did not influence daily nest predation rates for open- or closed-nest species. Although nest predation is high, at about 40%, incidence of predation events appears to be unpredictable – or even random.

pic snipe nest

In cold spring conditions, Icelandic Snipe are not able to hide their nests

Daily nest predation rates were significantly higher for closed nests (Redshank, Snipe and Black-tailed Godwit nests) in which a greater percentage of the clutch was visible. This suggests that the onset and rate of vegetation growth could potentially constrain the availability of suitable nesting locations for these species, and hence influence nest success, particularly among early season nests. This has been studied in Icelandic Black-tailed Godwits by José Alves and colleagues and is described in From local warming to range expansion.

For closed-nest species, the visibility of nests was significantly greater during the early part of the 2015 breeding season, when compared to 2016, due to slower grass growth in cooler conditions.  The higher predation rate of more visible nests of closed-nesting species was apparent even though nests were predated up to three weeks after egg visibility was measured. These findings suggest that early nesting attempts by concealed-nest species are unlikely to be successful in years when vegetation growth is delayed or slow. There can be major benefits of hatching early, with recruitment into breeding populations typically being lower for later-hatched chicks, so vegetation growth rates are likely to be really important to species that conceal their nests (Redshank, Snipe & Black-tailed Godwit in this study). However, given the ongoing trend for warmer springs at subarctic latitudes, the conditions in which early nests can only be poorly concealed are likely to be reducing in frequency.

In summary

pic goldie nest in habitat

Golden Plover nest set within a homogeneous habitat matrix

Perhaps surprisingly, nest predation rates were similar for open-nest and concealed-nest species and did not vary with vegetation structure in the surrounding landscape. However, nest-concealing species were about 10% more likely to have nests predated when the nests were poorly concealed, and the frequency of poorly concealed nests was higher at the start of the breeding season in colder conditions.

The paper at the heart of this blog is:

Vegetation structure influences predation rates of early nests in subarctic breeding waders. Rebecca A. Laidlaw, Tómas G. Gunnarsson, Verónica Méndez, Camilo Carneiro, Böðvar Þórisson, Adam Wentworth, Jennifer A. Gill and José A. Alves. IBIS. doi:10.1111/ibi.12827

pic sheep


GFA in Iceland

WaderTales blogs are written by Graham Appleton, to celebrate waders and wader research.  Many of the articles are based on previously published papers, with the aim of making wader science available to a broader audience.

@grahamfappleton

 

 

Which Icelandic Oystercatchers cross the Atlantic?

blog ringed birdMost Icelandic Oystercatchers leave Iceland in the autumn, crossing the Atlantic and arriving in Ireland, the UK and mainland Europe. Despite much shorter day-length and colder conditions, 30% spend the winter in Iceland, as discussed in this blog (Mission Impossible).

A 2020 paper from Verónica Méndez and colleagues asks whether birds that stay in Iceland or cross the Atlantic differ in sex, body-size or breeding location (within Iceland) and whether birds behave differently in different years. Interestingly, they were also able to test whether there is assortative mating – do Oystercatchers that stay in Iceland pair with other stay-at-home birds?

Iceland’s Oystercatchers

Iceland lies at the northern edge of the breeding range of Eurasian Oystercatchers. The country supports an unusually high proportion of wintering Oystercatchers, given its latitude and winter temperatures (Þórisson et al. 2018), and this may be influenced by the trans-oceanic flight of at least 700 km that migratory individuals must undertake to reach the European wintering sites. Individuals that migrate or stay within Iceland could differ in body size, for example if size influences the capacity to survive adverse winter conditions. Females tend to be slightly larger, and thus any sex differences in migratory behaviour could potentially reflect differences in body size.

It would be easy to imagine a scenario in which Oystercatcher pairs try to breed as early as possible, as this could increase the number of potential nesting attempts, as discussed in Time to nest again, based on Morrison et al. 2019. Does this mean that stay-at-home birds pair off at the start of the season, before migrants arrive? Given that most birds will choose the same mate in successive years, what happens in mixed pairs if a resident is waiting around for a delayed migrant? Is divorce likely to occur, as discussed in the importance of synchrony for Black-tailed Godwits, and could this mean that mixed pairs are rare?

Colour-marks and isotopes

blog map of sightingsA huge amount of the evidence that was used to answer the questions posed by Verónica and her colleagues was provided by volunteer birdwatchers, who reported colour-ringed Oystercatchers in their wintering areas, in the period through to April 2018. There’s an impressive set of dots on the map alongside, from the north of Scotland through to Spain, and the number of sightings continues to rise. It’s perhaps unsurprising, given the direction of travel from Iceland, that there is a strong westerly bias to the distribution across the British Isles.

The sample size from colour-ring observations was not big enough to answer all of the questions posed above, especially relating to whether pairs mate assortatively. To provide one data-point, it’s necessary to know the winter locations of both members of the pair. Might feather isotope ratios provide some help?

Oystercatchers that winter in Iceland use a restricted number of coastal sites (as inland sites are frozen during winter) and forage on marine prey. Elsewhere in Europe, a much wider range of marine and freshwater resources is available, with birds readily moving between the shoreline and fields, golf-courses, football pitches and road-side verges. Previous studies have shown that terrestrial diets produce different carbon and nitrogen isotope ratios, based on salinity and trophic level of prey items.  Are differences in habitat use and diet of Oystercatchers reflected in carbon (δ13C) and nitrogen (δ15N) isotope ratios of feathers grown during late winter?

blog isotope chest

Oystercatchers moult their chest feathers in late winter

The research team hoped that isotopic signatures from resighted colour-marked migrants and residents would be sufficiently different to be able to predict the probable wintering areas (Europe or Iceland) of marked birds not seen away from their breeding sites. If so, this would greatly increase the sample size, by enabling the combination of data from observations of colour-marked individuals with information on birds that could be assigned as Icelandic or European winterers using the isotopic composition of their feathers. They predicted that it would then be possible to:

  • Identify migratory strategies of individual Oystercatchers and explore whether the likelihood of migrating or staying in Iceland is related to gender, body size or breeding location.
  • Assess how consistent these individual strategies are between years.
  • Quantify spatial variation in the distribution of migrants and residents across the Icelandic breeding range.
  • Determine whether Oystercatchers mate assortatively in relation to migratory behaviour.

The work covered in this paper was conducted between 2013 and 2017. Full details of the study areas and methods are available in the paper – link below.

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Early nesting is not always a good idea – still incubating after sudden snow-fall

Migrant or resident?

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An Icelandic Oysterctaher spending the winter in Co. Sligo in Ireland

Of the 537 colour-ringed adults in the study, 58 were seen away from Iceland and 55 were shown to be resident in Iceland. Oystercatchers undertake a partial moult at the end of the winter period, when they grow new feathers on the neck and chest. The isotopic signature of a tiny piece of one of these feathers, taken from each adult at the time of ringing or recapture, was determined. The values of δ13C, which relates to habitat salinity, and δ15N, which relates to trophic level of diet, varied between residents and migrants but there was an overlap (details in paper).  There was enough of a difference, however, for it to be possible to allocate two-thirds of ringed birds that had not been seen in the winter period to the resident and migratory categories, with sufficient certainty, thereby increasing the sample size for other tests.

Consistency of migratory tendency

The 18 individuals that were observed in more than one winter were all consistent in migratory behaviour (10 residents and 8 migrants) and each was seen in the same specific location (Iceland or western Europe) in both winters. Where feather samples were taken in more than one year, there was no evidence of any bird changing its habitat or diet.

Factors influencing individual migratory programmes

Females and males were equally likely to migrate and there was no evidence that bigger (or smaller) birds were more likely to leave Iceland. Most Oystercatchers that winter in Iceland are in flocks in the west of the country, where the coast is warmed by the Gulf Stream. Unsurprisingly, westerly breeders were more likely to be resident than those in the south or northeast of Iceland (see figure below).

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Assortative mating

Vero and her colleagues were able to assign the migratory strategy to both members of the pair for 162 pairs (either by resightings or using predictions from isotopic signatures). Among these, 75 pairs (46%) were both migrants, 32 pairs (20%) were both residents and 55 pairs (34%) were mixed. The frequency of full-migrant, full-resident or mixed pairs varied significantly among regions, which was to be expected, given the differences in the likelihood of migrating from different parts of Iceland. There was no evidence of assortative mating; the likelihood of a particular individual pairing up with a migrant was as expected from the proportion of migrants in the area; it was not influenced by whether the particular individual was itself a migrant (see figure above).

To migrate or remain in Iceland

The consistency of adult migratory behaviour suggests that migratory strategy is determined in early life, and the regional variation in the frequency of migrants and residents may thus reflect variation in the conditions encountered by individuals during this life stage. As noted above, the frequency of residency is greatest amongst Oystercatchers breeding in the west of Iceland, which are the areas closest to the main wintering locations. Juveniles from the northwest and west are more likely to encounter these flocks of adult and sub-adult birds when moving south, than juveniles from the south, north-east and east, which are more likely to encounter migrating adults.

blog gen chThe regional variation in migratory strategy could arise through the influence of social cues, with juveniles adopting the behaviour of Oystercatchers they encounter and then recruiting back into their natal locations (more about this in Generational Change, focusing on Black-tailed Godwits). Birdwatchers across Europe will hopefully help to test this theory, by tracking colour-ringed juveniles during the early years of life. This is all part of a quest to identify the conditions that influence migratory behaviour and to understand the consequences, in terms of survival rates and productivity, of adopting different migratory traits.

Paper

Please click on the title below to access the paper:

Individual variation in migratory behavior in a sub-arctic partial migrant shorebird by Méndez V., Alves J.A., Þórisson, B., Marca, A., Gunnarsson, T.G., Gill, J.A.  Published in Behavioral Ecology (2020).  doi.org/10.1093/beheco/araa010 

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GFA in Iceland

WaderTales blogs are written by Graham Appleton, to celebrate waders and wader research.  Many of the articles are based on previously published papers, with the aim of making wader science available to a broader audience.

@grahamfappleton

 

Generational change

blog TGG on postIn a changing world, with more chaotic weather patterns and rapidly altering habitats, migratory birds are faced with opportunities and challenges. Long-term monitoring of colour-ringed Black-tailed Godwits, during a period of range expansion and phenological change, has revealed that individuals behave consistently over time but that the behaviour of new generations is moulded by the conditions they encounter.

A changing world 

When trying to explain observed changes in the distributions and annual cycles of migratory birds, there are many things to consider:

  • blog VM y flag

    Colour-rings enable life-time tracking. This bird, caught on its nest, had been ringed as a chick.

    Are individual birds able to take advantage of new breeding and non-breeding sites, as they become available, particularly if other areas become less suitable?

  • Are individuals able to change the timings and patterns of migration?
  • Do individuals adjust their migration routes as a consequence of changes in stop-over or wintering areas?
  • If individuals do not change what they do, how do we explain range expansions and changes in timing of migration?

Put simply, how does climate change lead to changes in distribution of migratory birds? Answering this question is key to being able to predict the rate and direction of future changes, and to assess whether our existing networks of protected sites will continue to support populations in the way that was intended. This issue was tackled by Jennifer Gill, José Alves and Tómas Gunnarsson in their paper “Mechanisms driving phenological and range change in migratory species”, published in Linking behaviour to dynamics of population and communities: applications of novel approaches in behavioural ecology and conservation, a special issue of Philosophical Transactions B (Royal Society).

Potential models

Change could happen in two main ways:

  • Individuals could relocate – having knowledge of a range of available conditions, they can choose to move elsewhere.
  • New generations could settle in new areas (in the breeding season, the non-breeding season or both) and/or adopt new migratory strategies.

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Map that illustrates range expansion

Working out whether change happens through individual movement or generational shifts can only be done by life-long tracking of individuals, in populations in which range change is happening. The Icelandic population of Black-tailed Godwit is ideal for such an investigation. Black-tailed Godwits have been expanding into new breeding areas of Iceland for over 100 years, as discussed in this WaderTales blog. Population growth has been facilitated through warming spring conditions, as discussed in From local warming to range expansion.

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Naive youngsters, gathering together before migration

Icelandic Black-tailed Godwits fly south in the autumn, to spend the winter in the British Isles, The Netherlands, Belgium, France, Spain and Portugal. As numbers have grown, winter counts have increased in many areas, with new flocks appearing and expanding on estuaries and areas of wet grassland where the species was previously absent or scarce.

 

Winter distribution

The Wetland Bird Survey shows that there are three times as many Black-tailed Godwits wintering in Great Britain as there were 25 years ago. The biggest changes in numbers have occurred on estuaries in the northwest of England, with the Morecambe Bay winter maximum rising from about 180 to 3200, for instance. Where have these extra birds come from?

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These young birds happen to have ended up on the Axe Estuary in Somerset

Black-tailed Godwits have been ringed in Iceland for nearly twenty years, providing a pool of known-age adults for which natal sites are known. Winter observations of colour-ringed individuals have shown an interesting pattern; birds breeding in newly-colonised areas, particularly in north and east Iceland, are the ones that are more likely to be found in newer winter sites.

In their paper, the authors suggest that birds nesting in these colder areas, where spring comes later, will be fledging quite late and leaving Iceland after adults have departed. With no experienced birds to follow, these young birds may well stop off at the first suitable site, many of which are in the north of the wintering range, and then they return to breed in their natal sites. Birds in Morecambe Bay don’t know that days are longer and the weather is kinder for other birds that travel further south to wintering areas such as Portugal.

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Wintering birds in Northwest England

Observations from birdwatchers show that the same colour-ringed individuals are nearly always found at the same wintering sites each year. Whatever mechanism is producing this new-breeding-site to new-wintering-site link, it is becoming clear that older birds continue to do what they have always done, with changes in distribution happening as a result of a generational shift.

The annual cycle

Colour-ringed Black-tailed Godwits have been tracked for over 25 years, with a small number of individuals contributing data for the whole of this period. This tracking information can be used to ask how much individuals move around and experience different sites and to assess whether individuals from different generations are using different parts of the range.

Using colour-rings, the Black-tailed Godwit team has discovered that, although individuals can live for over 20 years, in that time they generally use a total of only about four sites between leaving Iceland in late summer and returning in the spring. Basically, individual birds have very limited experience of sites and there is no evidence that they have moved to occupy different sites as, for instance, winter conditions have changed.

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Spring arrivals in Iceland

Colour-ring observations have shown that individual birds do not change their breeding or wintering locations and that migrating individuals often appear in the same stop-over sites year after year. The timing of movements is also pretty consistent, especially in the spring. A previous WaderTales blog called Why is spring migration getting earlier? demonstrated that the timing of  migration of individual Black-tailed Godwits varies very little, with observed shifts in the period of migration being driven by young birds returning to Iceland for the first time on average doing so somewhat earlier than previous generations. Once individual birds settle into a timing pattern, they stick to it.

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Black-tailed Godwits, newly arrived in Iceland after crossing the Atlantic

Migration patterns

As discussed above, individual Icelandic Black-tailed Godwits have experience of only a small number of sites, which they use on an annual cycle. When migrating, a bird will generally use the same stop-over site when breaking its journey south, to undertake autumn moult, or on their way north, to take on fat for the trans-Atlantic journey. There is a range of spring migratory strategies in islandica Black-tailed Godwits, as discussed in Overtaking on migration.

blog wwrwOnce established, the annual migratory programmes of individuals rarely change, as illustrated by the map to the right. Colour-ringed Black-tailed Godwit W-WR/W regularly moulted on the Wash, in eastern England, before spending the late winter and spring in northwest England. In the late summer of 2002 he was reported at Slimbridge on 18th and 20th July but back on the Wash on the 25th. Having made the Atlantic crossing and ended up in southwest England, he was able to correct what he may have perceived to be his mistake, returning to the moulting area that he had been using since at least 1996.

Individuals might not change their annual migration routes but we do see changes in numbers on different sites that are used during migratory stop-overs. In a paper published in 2018, Mo Verhoeven and colleagues investigated whether observed changes in migratory patterns of a population of the limosa subspecies of Black-tailed Godwit were caused by individuals altering their strategies or by generational change.

Limosa Black-tailed Godwits leave breeding areas in countries such as The Netherlands in late summer, heading south to either West Africa or Iberia, where they spend the winter. In spring they all gather in staging sites in Portugal and Spain, typically on rice fields. Over the course of less than ten years, the average peak number in Extremadura (Spain) has dropped from about 24,000 to 10,000, while the numbers on the Tagus and Sado estuaries rose from 44,000 to 51,000. These changes took place during a period of rapid population decline, as described in this blog focusing on a paper by Rosemarie Kentie and colleagues.

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Limosa Black-tailed Godwits feeding in a rice field in the Tagus estuary

Mo Verhoeven et al have shown that this rapid population-level shift in spring stop-over sites from Spain to Portugal, 300 km further west, was driven by young godwits increasingly using Portugal in the period January to March, instead of Spain. Nearly all of the older birds stuck with the routes they knew. The paper is Generational shift in spring staging site use by a long-distance migratory bird.

Change happens to birds

One thing that is becoming clear in Black-tailed Godwits is that birds are being affected by change – individuals do not have the knowledge or flexibility to effect change. Even in long-lived birds, like Black-tailed Godwits, we see no evidence of individuals altering what they do over what is now two decades, despite the fact that the species’ migration dates, wintering areas and migration routes have all perceptibly changed over the same time period. It’s all about generational change. The behaviour patterns of young birds arise from the conditions they encounter in the first year of life, after which they are repeated.

Details of the Generational Change paper by Gill et al

blog LJ sum plumThe paper at the heart of this blog is: Mechanisms driving phenological and range change in migratory species by Jennifer Gill, José Alves and Tómas Gunnarsson, from the Universities of East Anglia (UK), Aveiro (Portugal) and Iceland. It is published in Linking behaviour to dynamics of population and communities: applications of novel approaches in behavioural ecology and conservation, a special issue of Philosophical Transactions B (Royal Society).

The paper could not have been produced without the help of “thousands of observers of colour-ringed godwits who have made these analyses possible”. This WaderTales blog is a celebration of the work they do: Godwits and Godwiteers.


GFA in Iceland

WaderTales blogs are written by Graham Appleton, to celebrate waders and wader research.  Many of the articles are based on previously published papers, with the aim of making wader science available to a broader audience.

@grahamfappleton

 

Whimbrel: time to leave

blog WW-WLGeolocators* have provided fantastic information about the movements of migratory birds – making links between countries, revealing previously unknown stop-over sites and indicating just how quickly birds traverse our planet. A small number of Icelandic Whimbrel have carried geolocators for up to six annual cycles, providing Camilo Carneiro with an opportunity to investigate the annual consistency of egg-laying, autumn departure, arrival in West Africa, departure in the spring, stopover in Western Europe and arrival back in Iceland.

* Geolocators are tiny devices that record the daily positions of birds, by measuring the timing of dawn and dusk. An individual typically carries a geolocator for a year and then needs to be re-caught for the data to be downloaded.

Planning a trip

When booking a train journey on-line, the first question I am asked is whether I want to stipulate departure time or arrival time.  In early spring, with breeding on their minds, you might think that Whimbrel will focus on the time they need to be in Iceland, rather than the time they leave West Africa? If that’s the case then it might be best to take early spring opportunities if they arise, to catch express winds that will make the journey as rapid as possible and to get to Iceland early. Is that the case?

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The Whimbrel is one of several wader species that breed in Iceland. Each autumn, Redshank, Snipe, Golden Plover, Oystercatcher and Black-tailed Godwit fly south to Europe, especially Ireland and the United Kingdom, but many Ringed Plover, most Dunlin and most Whimbrel travel as far as Africa. The main wintering sites for Whimbrel are in West Africa, south of the Sahara, in countries such as Guinea-Bissau. Here they can be seen feeding on crabs on the mangrove-fringed muddy shoreline (above). It’s a very different environment to the inland floodplains of Iceland (below).

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In a paper by Tómas Gunnarsson & Gunnar Tómasson in 2011, we learnt that Whimbrel arrival times in Iceland did not change much between 1988 and 2009 (just 0.16 days earlier per year), while timing of arrival was advancing much more in species that travel less far to winter grounds, as you can see in this diagram.

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Changes in first spring arrival dates of six species of waders in southern Iceland from 1988 to 2009 (reproduced from Gunnarsson & Tómasson 2011).

The arrival date for Black-tailed Godwit was advancing fastest (0.81 days per year). In more recent research, it has been shown that the rapidly advancing trend for Black-tailed Godwits is being driven by new recruits to the population – individual adults are not changing their schedules. Why is spring migration getting earlier? summarises a paper by Gill et al in Proceedings of the Royal Society B.

The Whimbrel trend has been recalculated, with a longer run of years, and the new change of 0.03 days earlier per year is not significantly different from zero. Given that Whimbrel are breeding alongside other species that are arriving in Iceland much earlier than thirty years ago, what are the constraints to the timing of their migrations?

Migration timings for Whimbrel

Camilo Carneiro, José Alves and Tómas Gunnarsson from the Universities of Aveiro (Portugal) and Iceland have been studying a population of Whimbrel in Southern Iceland. Birds are caught on the nest in one year and then re-caught in the subsequent year – or two years later if a bird evades capture in the intervening summer. The following results summarise weeks and weeks of patient fieldwork and brush over the hours of frustration caused by wary birds that have been caught before!

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Over the course of the whole study, 86 Whimbrel were fitted with geolocators, 62 of which were retrieved. Repeatability could be calculated for 16 birds, with between 2 and 7 years of data collected from each individual. The results are summarised in these few bullet points. Please see the paper for confidence intervals and more details about differences between the sexes.

  • Blog tagIndividual timings of autumn departure from Iceland varied between years. The repeatability index is 0.28, with a suggestion of a gender difference (females 0.40, males 0.02). Males tend to look after chicks for a longer period than females so their departure dates may be more strongly influenced by the success of each year’s breeding attempt.
  • Autumn arrival time in West Africa was closely linked to departure time because, on all but one occasion, southward migration was achieved through a single direct flight. See Iceland to Africa non-stop.
  • Spring departure time from West Africa was highly consistent, with a repeatability index of 0.76 and no discernible difference in repeatability between males and females.
  • blog long green grassSpring arrivals in Iceland. Some Whimbrel that managed to complete spring migration in a single flight in some years stopped off in other years. These breaks, perhaps to wait for more helpful wind conditions and/or to refuel, resulted in variability in annual arrival dates for individuals. The repeatability for the two sexes combined was 0.23.
  • Laying date was the least consistent stage of the annual cycle, with a repeatability index of 0.11 and no significant difference between males and females.

In an individual Whimbrel’s annual cycle, there appears to be one fixed point – departure from wintering ground in West Africa. With no discernible seasonality of resource availability on the wintering grounds and little change in day length in these areas, departure dates are probably being determined by an ‘internal clock’. Two major unknowns will then determine what happens in the next twelve months. Will wind and weather conditions be conducive to a one-leg flight to Iceland and how successful will a bird be in any particular breeding season? Unforeseen events, such as having to wait for a delayed partner, losing a first clutch, and the time needed to guard chicks will all affect the timing of autumn migration.

Understanding individuals

blog tag through grassThe study of wader migration has advanced hugely.

  • Fifty years ago, the main measure of migration phenology was the appearance of the first individuals of a species.
  • Colour-ring sightings are ideal for providing repeat arrival dates over the lifetimes of individuals, as exemplified by the Gill et al paper on Black-tailed Godwits, which suggest that individual timing is highly repeatable.
  • Geolocators have provided more detailed information about the precise arrival and departure timings of individuals, which is so important if we wish to conserve threatened, migratory species that visit areas in which data collection was previously virtually impossible.
  • Now, by tracking individual birds for several years, it is possible to look at the variability in annual patterns, and what can cause this variability.

Over the next decade or so, as devices get smaller and remote downloads become easier (eg using GSM tags), it should become possible to understand the conditions that lead to fast, slow and aborted migratory journeys in a whole range of species. Exciting times!

Paper

Why are Whimbrels not advancing their arrival dates into Iceland? Exploring seasonal and sex-specific variation in consistency of individual timing during the annual cycle. Camilo Carneiro, Tómas G Gunnarsson & José A Alves. Frontiers in Ecology & Evolution.

There is more about the information that is obtained from geolocators, how they work and the affects that they have on the individual birds that wear them in these two blogs:

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GFA in Iceland

WaderTales blogs are written by Graham Appleton, to celebrate waders and wader research.  Many of the articles are based on previously published papers, with the aim of making wader science available to a broader audience.

@grahamfappleton