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 12^th 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

• The edge effect of forests is discussed in work from Estonia by Triin Kaasiku (see Keep away from the trees). Here she looks at the outcomes of nests at different distances from forest edges.
• Do waders avoid nesting close to trees? This is discussed in Mastering Lapwing Conservation.
• The predator effect lingers long after forests are cleared, according to research described in Trees, predators and breeding waders.

Work by Aldís Pálsdóttir (pictured right)

• The WaderTales blog, Power lines and breeding waders, summarises Effects of overhead power-lines on the density of ground-nesting birds in open sub-arctic habitats.
• Further blogs about wader numbers close to roads and in the vicinity of summer cottages will be added when papers are published.

Changing agricultural systems in Iceland (work by Lilja Jóhannesdóttir)

• Do Iceland’s farmers care about wader conservation? At a time of agricultural expansion, are farmers prepared to leave space for breeding waders?
• Farming for waders in Iceland investigates densities of breeding waders along the gradient of agricultural intensification associated with farming activities.
• Designing wader landscapes investigates whether breeding densities of waders can be maintained, as farming expands and intensification increases.

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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.

Power-lines and breeding waders

Around the globe, people are consuming more energy, much of which is delivered to cities, towns, individual homes and businesses via over-head electricity cables. In a paper in Ibis, Aldís E Pálsdóttir and colleagues investigate the effects of power-lines on Iceland’s breeding waders. This is the first of several papers from Aldis’ PhD thesis, in which she seeks to understand how forestry and the sprawl of new infrastructures (roads, cottages and power-lines) are changing bird distributions within what were previously open landscapes.

Breeding waders in Iceland

Iceland is a hot-spot for breeding waders, holding half or more of Europe’s Dunlin, Golden Plover and Whimbrel, in a country that is a bit smaller then England. In a 2017 report prepared by AEWA (Agreement on the Conservation of African-Eurasian Migratory Waterbirds), in response to concerns about the effects of afforestation on Iceland’s waterbirds, we learn that:

“Iceland is second only to Russia in its importance as a breeding ground for migratory waterbirds in the AEWA region. It supports the most important breeding populations in Europe for six species of waders and is the second most important country for three.”

Data in the table alongside have been extracted from Annex 4 of the report, which was discussed at the 12^th Standing Committee of AEWA in Jan/Feb 2017.

Power lines

For waders nesting in Iceland, power-lines are a new addition to a once-open landscape. Pylons provide potential nesting opportunities for Ravens and perches for Gyrfalcons, while the wires between them are a collision risk. Under power-lines, carcases of swans, geese and waders may attract scavenging Arctic Foxes and Ravens, thereby increasing the activities of nest predators. Perhaps these actual and perceived threats affect densities of breeding waders in the vicinity? Or might birds react to something less obvious, such as the emission of UV light or electromagnetic radiation?

In Iceland, the vast majority of electricity is produced from hydropower or geothermal sources, often long distances from the areas in which the power is used. A new move to develop the wind energy sector has the potential to further add to the number of power lines and introduce them in more areas of the country. Much of the increase in electricity production over the last fifty years has been used to fuel industries such as aluminium smelting and there is the potential to further expand generation capacity, perhaps exporting some electricity to other countries.

Much of Iceland’s electricity is used to power aluminium smelters (here, in Hvalfjörður)

Counting the birds

Aldís counting waders on a transect

Aldís conducted the fieldwork for this study between the 6^th May and the 20^th June 2019, counting birds along 85 transects of between 300 m and 500 m, running perpendicular to power lines. The full methods are described in the paper but it is interesting to see that they included a check to see whether there were different results if walking towards or away from the power lines. Each transect was divided into intervals of 50 m length, each corresponding to 1 hectare of surveyed land. For each power line, Aldís recorded the number of cables, pylon characteristics, and the height and voltage of the line.

Results

In total, 1067 birds of 21 different species were recorded on the 85 transect surveys. Over 90% of sightings were of eight species considered in the subsequent analysis: Dunlin, Black-tailed Godwit, Golden Plover, Meadow Pipit, Redshank, Redwing, Snipe and Whimbrel. Having analysed the data, Aldís and her colleagues concluded that:

• For all eight species combined, the areas closest to the power lines (0-50 m) supported densities of approximately 112 birds/km² (±13 SE) which increased by approximately 58% to 177 birds/km² (±24 SE), in the sector that was between 450-500 m away from the power-lines. On average, there was a 4% increase in abundance between adjacent 50 metre bands.
• At the species level, Redshank (figure below) and Whimbrel density increased significantly with distance from power lines (18% and 9% per 50 m, respectively) but no other significant effects were detected for other species individually.
• There were no detectable difference between types of power-lines or relating to the voltages of the electricity they carried.

Implications of the research

In the paper’s discussion there are questions as to why densities of Redshank and Whimbrel (right), in particular, are lower near power-lines. The two species behave differently while nesting, with Redshanks being nest-hiders and Whimbrel nesting in the open, but previous research has shown that their nest predation rates are quite similar (see Where to nest?).

The reason why significant reductions in density close to power lines were apparent for Whimbrels and Redshanks (but not for other species) is not clear but the authors suggest that sample sizes may have been too low for there to have been measurable effects for species such as Dunlin and Golden Plover (below).

Power lines could have direct impacts, such as increased collision risk, but this may be difficult to establish directly, as the authors suggest that carcasses are likely to be quickly removed by scavengers.

Ravens may find it easier to find and predate nests if there are pylons or wires on which to perch but it will be hard to discriminate between an actual predation effect, reducing numbers in areas close to power-lines, and the avoidance of risky areas because of a perceived threat of predation. This is discussed in Mastering Lapwing conservation.

Given the depressed density of ground-nesting bird species in the vicinity of overhead power lines, the authors of the paper suggest that burying power lines might be a better option, even though there would be temporary disturbance to the ground during installation.

What are the implications for Iceland’s breeding waders?

It would be interesting to calculate how many Whimbrel and Redshank (left) territories would be lost over the course of a 50 km run of power-lines through open landscapes – and then extrapolate that to 500 km and 5,000 km. As shown in the earlier table, 75% of Europe’s Whimbrel breed in Iceland. How vulnerable are they to power-line infrastructures and what might be the impact on a breeding population of over 300,000 pairs?

This is the first of several papers from Aldis’ thesis, in which she seeks to understand the current rapid changes to Iceland’s lowland landscapes. Links to other blogs and papers will be added as they appear. It should soon be possible to reveal the combined effects of these incursions into open wader habitats, by considering plans that might affect these areas over the next twenty years, working out potential losses and setting these numbers in a flyway context.

This paper is published as:

Effects of overhead power-lines on the density of ground-nesting birds in open sub-arctic habitats. ALDÍS ERNA PÁLSDÓTTIR, JENNIFER A. GILL, SNÆBJÖRN PÁLSSON, JOSÉ A. ALVES, VERÓNICA MÉNDEZ, BÖÐVAR ÞÓRISSON & TÓMAS G. GUNNARSSON. Ibis. https://doi.org/10.1111/ibi.13089

Here’s a link to another blog about Aldís Pálsdóttir’s research: Iceland’s waders need a strategic forestry plan.

A complementary set of papers by Lilja Jóhannesdóttir investigated how changes to Iceland’s farming may also be affecting breeding waders. These are discussed in three WaderTales blogs:

• Do Iceland’s farmers care about wader conservation? At a time of agricultural expansion, are farmers prepared to leave space for breeding waders?
• Farming for waders in Iceland investigates densities of breeding waders along the gradient of agricultural intensification associated with farming activities.
• Designing wader landscapes investigates whether breeding densities of waders can be maintained, as farming expands and intensification increases.

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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.

Where to nest?

There 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

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.

Iceland: a wader factory

As 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).

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.

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.).

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.

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.

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

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

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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

 

 

Fennoscandian wader factory

 

Nesting Temminck’s Stint – the smallest of the 22 wader species for which trends are reported

At the end of the summer, vast numbers of waders leave Norway, Sweden and Finland, heading southwest, south and south-east for the winter. In a 2019 paper by Lindström et al, we learn what is happening to these populations of Fennoscandian breeding species, as diverse as Temminck’s Stint and Curlew. The news for the period 2006 through to 2018 is basically pretty good – most populations have been stable and there are even some that have increased – but there are worrying signs for Broad-billed Sandpiper, Red-necked Phalarope and Whimbrel.

Breeding waders of Fennoscandia

As a volunteer taking part in the Breeding Bird Survey (BTO/JNCC/RSPB) in the UK, I feel that I do my bit to monitor what is happening to local bird population – providing counts that build into national trends. The work involved in delivering indices for breeding waders across the area of Fennoscandia shown in the map is in a different league. Here, counters visit habitats as diverse as forests, wetlands, mires and tundra, within the boreal and arctic areas of Norway, Sweden and Finland. Some survey sites are so remote that access requires the use of helicopters.

Fennoscandia provides important breeding areas for a large set of wader species, and models suggest that these habitats may be particularly vulnerable to climate change, especially increasing summer temperatures. The 2006-18 analysis in Wader Study, the journal of the International Wader Study Group, presents population trends for 22 wader species. The trends are based on 1,505 unique routes (6–8 km long), distributed over an area that’s about four times that of the United Kingdom. 

The surveys took place across the whole of Norway and Finland, and in the northern two thirds of Sweden, between 58°N and 71°N, which largely coincides with the boreal, montane and arctic regions of Fennoscandia. The systematic distribution of these routes ensures that the main habitats in these countries are sampled in proportion to the area they cover. The paper describes the methodologies used in the three countries and the way that data were combined, especially factors used to translate sightings of individuals into ‘pair-equivalents’.

Overview of results

Looking at the results from across Norway, Sweden & Finland:

• In terms of pure numbers, Golden Plover was the most commonly encountered wader species, followed by Wood Sandpiper, Snipe, Greenshank and Green Sandpiper.
• The five most widespread species, seen on the highest number of routes, were Snipe, Green Sandpiper, Greenshank, Wood Sandpiper and Common Sandpiper.
• Wader species richness and the total number of wader pairs were both higher with increasing latitude; the median number of wader pairs per 10 km increased from just below 3 at latitudes 58–60°N, to just above 26 at latitudes 69–71°N.
• Using a multi-species indicator, the research team found no general change in wader numbers over the period 2006-18.
• The trends were significantly negative for three species: Red-necked Phalarope (-7.9% per year), Broad-billed Sandpiper (-5.4% per year) and Whimbrel (-1.3% per year).
• The trends were significantly positive for three species: Oystercatcher (+4.9% per year), Dunlin (+4.2% per year) and Wood Sandpiper (+0.8% per year).
• There was no significant trend for another 16 species for which encounters were deemed to be frequent enough for analysis.
• Population trends of long-distance migrants tended to be more negative than those of medium-distance migrants. This is discussed in detail in the paper.

Focusing on some key species

The Lindström et al paper is a tremendously rich source of information and references. Here are some species-specific highlights.

Oystercatcher. In the context of a species that is declining across NW Europe, the fact that there is a significant increase in Oystercatchers across Fennoscandia may be surprising. However, the authors note that there was a jump in numbers between 2006 and 2007 with little change since then.

Lapwing. The trends within the three Fennoscandian countries are very different. In Norway, there has been a dramatic decline (-15.2% per year during 2006–2018) and the Lapwing is now nearly extinct in many areas. The trend in Sweden is also significantly negative (-5.8% per year). In Finland, however, where the species is more widespread and numerous, there has been a strong increase (+5.9% per year) during the same period. See figure alongside.

Golden Plover. No significant change overall. There are some country-specific differences in trends, with a moderate decline in Norway being countered by a moderate increase in Sweden. 

Snipe. The overall trend of this species for each country indicates an initial decline followed by an increase. A similar pattern has been noted in the UK’s Breeding Bird Survey over the same period. 

Nesting Whimbrel

Woodcock. The trend for 2006–2018 is basically stable and similar in all three countries.

Curlew. There is no significant trend, overall, but populations in Norway and Sweden have declined at the same time that numbers in Finland have increased.

Whimbrel. Fennoscandian trend indicates a decline of 1.3 % per year. Whimbrel is doing poorly in Norway and Sweden but better in Finland. 

Wood Sandpiper. This widespread species has increased slowly (0.8% per year), a trend that is largely driven by Norwegian and Swedish populations.

Wood Sandpiper was the second most commonly encountered wader

Redshank. The fact that no change was discernible, suggests that boreal and arctic populations are faring much better than the breeding populations further south in Europe. For example, see Redshank – warden of the marsh.

Redshank – more obvious than most breeding waders encountered!

Spotted Redshank. The estimated annual decline for Spotted Redshank is 2.8% per year but the species is too thinly spread for this to provide significant evidence of a decline. This rate is very similar to the recent drop in the Wetland Bird Survey index in the UK. See Fewer Spotted Redshanks.

Broad-billed Sandpiper. This species has the second most negative trend among the 22 species analysed (-5.6% per year). The bulk of information comes from Finland where the trend is even more negative (-7.5% per year). Birds head southeast in the autumn to countries bordering the Indian Ocean – areas for which winter trend data are not available. The species is still considered to be of ‘least concern’ but perhaps this designation may need to be revisited?

Dunlin. Breeding birds in the survey area are largely of the alpina race. The overall trend is significantly positive (+4.1% per year), which is in sharp contrast to the strong declines of the schinzii subspecies that breeds around the Baltic Sea, western Finland and further south and west in Europe.

Ruff. There were major declines in the period immediately prior to this review (Lindström et al. 2015) but changes reported here are lower (-2.3% per year) and the decline is not statistically significant.

Red-necked Phalarope. The authors write, “This species has the most negative trend of all the 22 species [-7.9% per year], with most data coming from Sweden. We do not know the cause of this decline but, given that this species shares its south-eastern migration route with Broad-billed Sandpiper, whose population exhibits the second largest decline, the relevant problems might largely apply somewhere along the migration routes”.

Link to Britain & Ireland

As shown in Which wader when and why? there are strong migratory connections between Fennoscandia and the British Isles. Some waders, such as Green, Common and Wood Sandpipers, pass through on their way south in the autumn, whilst many more fly here for the winter, to take advantage of the warmer maritime climate.

Three wader species with particularly strong links between Fennoscandia and Britain & Ireland are still shot and eaten in these islands. Each autumn, large numbers of Woodcock, Golden Plover and Snipe cross the North Sea. It is difficult to ascertain figures for the number that are shot but there is agreement that the vast majority are winter visitors, as opposed to native birds. The results presented in the paper suggest that there have been no discernible changes in the Fennoscandian populations of these three game species in the period 2006-18. Two earlier WaderTales blogs focus on Woodcock and Snipe in Britain & Ireland:

• Conserving British-breeding Woodcock
• Snipe & Jack Snipe in the UK & Ireland

There has been no significant change in Golden Plover numbers across Fennoscandia

Two WaderTales blogs about wintering waders in Great Britain and the island of Ireland were published in 2019, based on reviews in British Birds and Irish Birds. These were Do population estimates matter? and Ireland’s wintering waders. The six big losers, in terms of wintering numbers in these islands, were Knot, Oystercatcher, Redshank, Curlew, Grey Plover and Dunlin. Knot arrive from Greenland and Canada, with Grey Plover flying from Russia, but it is interesting to think about this Fennoscandian breeding analysis in the context of winter losses of the other four species.

• Wintering numbers of Oystercatchers have dropped recently in Britain and in Ireland. The population is made up of migrants from Iceland (more about this here), very large numbers from Norway, birds that stay within the British Isles and smaller numbers from other European and Scandinavian countries. Given there is no discernible decline in Fennoscandia, it seems likely that much of the decline can be attributed to a major fall in Scottish breeding numbers (more about this here).
• Most Redshank wintering in Britain & Ireland are of local or Icelandic origin. Fennoscandian numbers seem to be stable; if there were any changes, these would probably not be apparent in wintering numbers within the British Isles.
• The Eurasian Curlew has been classified as ‘near-threatened’ and the species is known to be declining in many areas (see this blog about serious problems in Ireland). Ringing shows a particularly strong link between Finland, where breeding numbers seem to be increasing, and Britain & Ireland. The decline in British and Irish winter numbers is probably being driven by lower breeding numbers within the British Isles and in countries such as Sweden, Norway and Poland.
• There is a theory that new generations of alpina Dunlin may be more likely to winter within Europe’s mainland estuaries, instead of continuing their westward migration across the North Sea. This might explain the apparent anomaly between the 4.1% per annum rise in Fennoscandian numbers and recent winter declines of 3% in Britain and over 20% in Ireland.

Going forwards

Some of the survey areas were in particularly remote areas

Many of the study squares that were covered during these surveys are a long way from the main centres of human population in Norway, Sweden and Finland. The governments of the three countries are to be congratulated for supporting this important monitoring, which relied on the commitment of hundreds of volunteers. It is to be hoped that these surveys will continue and that further species-focused work will be able to explain some of the differences across Fennoscandia, particularly between eastern and western areas. The rapid declines in numbers of two species that migrate southeast each autumn (Broad-billed Sandpiper and Red-necked Phalarope) highlights the need for better information about what is happening on the flyway linking Fennoscandia with the Arabian Sea and coastal countries of the Indian Ocean.

Paper

Population trends of waders on their boreal and arctic breeding grounds in northern Europe: Åke Lindström, Martin Green, Magne Husby, John Atle Kålås, Aleksi Lehikoinen & Martin Stjernman. Wader Study 26(3)

Click on the title of paper to access it on the International Wader Study Group website. Paper is only available to members of IWSG. If you have read the whole of this blog you’ll probably want to join!

Nesting Bar-tailed Godwit in smart summer plumage

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Graham (@grahamfappleton) has studied waders for over 40 years and is currently involved in wader research in the UK and in Iceland.  He was Director of Communications at The British Trust for Ornithology until 2013 and is now a freelance writer and broadcaster.

 

 

January to June 2019

One or two WaderTales blogs are published each month. The series is UK-based with a global reach. Suggestions of newly-published research on waders that might be of interest to birdwatchers who appreciate waders/shorebirds are welcomed. I am particularly keen to give feedback to colour-ring readers; they provide a huge amount of information that lies at the heart of these stories.

Click on a link in bold to read an individual blog.

• Winter territories of Green Sandpipers includes unpublished information from southern England, where survival is affected by the severity of winters.
• In search of Steppe Whimbrel summarises a paper about two very special individual Whimbrel. Will this knowledge help to rescue a subspecies?
• From local warming to range expansion explores the role of climate warming in fuelling the century-long range expansion of Iceland’s Black-tailed Godwit/
• Ireland’s Curlew Crisis focuses on the nationwide breeding survey between 2015 and 2017, which revealed a 96% decline in the number of pairs in just 30 years.
• Do population estimates matter? is inspired by the waders section of Population estimates of wintering waterbirds in Great Britain, based on data from the Wetland Bird Survey and the Non-estuarine Waterbirds Survey.
• The Waders of Northern Ireland was written as a promotional tool for a 2019 breeding survey but covers wintering and passage species too.
• Ireland’s wintering waders provides the latest population estimates of waders wintering in Ireland, set within in a European context.
• Redshank – the ‘warden of the marsh’ focuses on Redshank that breed on saltmarshes and the agricultural subsidies that help to fund their conservation.
• Not-so-Common Sandpipers mixes information about migration with a review of Common & Spotted Sandpipers by Phil Holland.
• Chicks and Ticks reviews a study of the effects of ticks on the survival probability of Golden Plover chicks.
• Managing water for waders celebrates work to reduce flooding, store fresh water for farmers and create habitat for breeding waders.

You can sign up to receive an e-mail notification when a new blog is published.

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Graham (@grahamfappleton) has studied waders for over 40 years and is currently involved in wader research in the UK and in Iceland.  He was Director of Communications at The British Trust for Ornithology until 2013 and is now a freelance writer and broadcaster.

Chicks and ticks

How do ticks affect Golden Plover chicks? By utilising data from an existing study, David Douglas and James Pearce-Higgins have discovered that Golden Plover chicks that carry more sheep-ticks Ixodes ricinus have a lower chance of survival. Their findings are written up as a paper in Bird Study. The work is only based on a small sample and the data don’t identify the mechanism that leads to increased mortality but, given the current interest in the biological effects of ticks, the findings are interesting.

Costs of carrying ticks

Carrying ticks has three potential effects on wader chicks

• Ticks suck blood, which could be costly.
• Ticks can introduce diseases, via tick-borne bacteria and viruses.
• There may be effects on feeding efficiency, via impaired vision, hearing etc.

Sheep ticks act as vectors for a variety of pathogens, including the louping ill virus (LIV) which can affect a range of domestic and wild mammals, as well as wild birds. LIV is known to cause high mortality of Red Grouse chicks but there has been no previous assessment of the effects of sheep ticks on other moorland birds, such as breeding waders. It should be noted that wader chicks eat ticks – so they are not ‘all bad’.

Ticks can clearly be seen on the unfeathered lower eyelid of this young chick. There are smaller ticks attached to the gape at the base of the beak as well.

All sorts of things could influence the probability of tick infestation in birds:

• Ticks may be able to survive better in warmer conditions.
• Tick numbers can be affected by the number of mammalian hosts.
• Mammal to bird transfer could be affected by land management and habitat structure.

Ecological interactions between ectoparasitic ticks and waders are not well understood. Given possible increases in tick abundance with climate change, the authors of the new study felt that it would be useful to test whether ticks have detectable effects on the Golden Plover chicks that carry them.

Spotting an opportunity

The Golden Plover chicks that provided the data used in this paper were caught as part of a wind farm study at Gordonbush in northern Scotland, a site made up of 33 km² of blanket bog. There is more information about the study in these two papers:

• Negative impact of wind energy development on a breeding shorebird assessed with a BACI study design. Alex Sansom, James W. Pearce‐Higgins & David J. T. Douglas
• Relative importance of prey abundance and habitat structure as drivers of shorebird breeding success and abundance. David J. T. Douglas & James W. Pearce‐Higgins

Gordonbush, prior to the development of a wind farm

Tick numbers on Golden Plover chicks were collected at the time of ringing (within 24 hours of hatching) and during subsequent recaptures. Recapture was facilitated by locating tagged birds using radio-location. On each capture, chicks were weighed and the number of ticks visible on the bare parts of the head (around the eyes and bill) were counted. Most ticks attach themselves to the bare parts of the head and neck.

Variation in tick loads on Golden Plover chicks

Incubating adult. Golden Plovers commence primary moult at the start of (or before) incubation. Read more here.

The number of sheep-ticks found on these Golden Plover chicks was higher than those previously reported for waders but were within the range of those found on Red Grouse on moorland. Previous wader studies had been focused upon areas with sheep, which were routinely treated to reduce tick infestations. In the current study it was found that:

• 90% of chicks were carrying ticks, with between 1 and 12 ticks being found on each affected Golden Plover chick. The highest tick-load was found in mid-age chicks.
• Tick loads were higher during periods with warmer maximum temperatures and when chicks were estimated to have moved through taller vegetation between recaptures.
• Chick growth rates were depressed by high tick-loads, especially when temperatures were warmer.
• Of the 21 chicks, 4 fledged, 13 died and the outcomes for the other 4 were unknown. Half of the deaths appeared to have been due to predation and half to starvation/exposure.
• Chicks that were heavier (for their ages) were more likely to survive. Those with higher tick loads (for their ages) were less likely to survive.

With the small sample size, it was not possible to detect a correlation between tick load and chick growth rates but low survival was correlated with high tick-loads. This had not previously been documented for waders.

Implications for wader conservation

Gordonbush is an area where there is no grazing by domestic animals so the likely mammalian tick-hosts are Red Deer in particular and also Mountain Hare. The correlation between warmer weather and tick numbers, found in this study, could be explained by increased tick activity, while the link to taller vegetation may well be explained by ticks seeking damper microhabitats. In their discussion of the results, the authors suggest potential ways that ticks and waders, of different ages, might interact. Anyone looking to expand the work, in order to understand the mechanics of tick infestation, is likely to spend more time looking at ticks than waders!

Could ticks be reducing survival probabilities for  young Curlew?

The authors of this paper were not able to test for the presence of disease (such as LIV) in their Golden Plover population but this is a plausible cause of the increased probability of mortality. In a study in Yorkshire by Newborn et al. (2009), no evidence was found of LIV in wader chicks, whereas it was present in 3.6% of a sample of Red Grouse chicks at the same sites. Newborn and colleagues report that a single Eurasian Curlew chick has previously been recorded to be seropositive for LIV. In the Newborn study, the lowest incidence of ticks among waders was in Lapwings (6% of broods), followed by Golden Plover (47% of broods had ticks) and Curlew (91% of broods). There is a hint, in these data, that ticks might more commonly attach themselves to wader chicks that are found in taller vegetation.

Despite high tick loads on chicks, and the correlation with lower chick survival, the overall percentage of Golden Plover chicks known to fledge in the Gordonbush study (19%) is comparable with other studies. Perhaps ticks are only causing the deaths of chicks that would have died anyway?

The authors suggest that no case should be made for tick-control, to help breeding waders, until it is clear whether tick-based chick mortality limits Golden Plover and other wader populations on moorland. In the paper, they argue that previous attempts to reduce moorland tick abundance and tick loads on Red Grouse have failed to detect convincing evidence of improvements in grouse survival, breeding success or post-breeding densities.

A range of methods have been deployed in attempts to reduce moorland tick abundance and tick loads on Red Grouse, including reducing densities of mammalian tick hosts (Mountain Hare and deer) and deploying acaricides (as used in sheep dips). See this link to material from GWCT on ticks and Red Grouse.

In conclusion

Work at a single site over two years appears to have documented a level of tick infestation in Golden Plovers that is associated with chick mortality. It is not clear how chicks are being affected, particularly given that there is insufficient evidence thus far that ticks affect chick growth rate.

The authors collected the data analysed in this paper for other studies – the focus was not on tick effects – and they hope that funding might be found for future research focusing upon the associations between ticks and waders, other birds and other animals. Until that happens, it would be useful if shorebird biologists who repeatedly handle wader chicks, in order to measure growth and survival rates, could routinely record the presence or absence of ticks.

Given that warming temperatures could lead to increased tick abundance, this seems to be a good time to discover more about tick behaviour, the importance of ticks as a food source for wader chicks and whether tick-loads are reducing growth rates and fledging success in other wader species.

Paper

This research is published in the BTO journal Bird Study. Click on the details below to link to the full paper:

Variation in ectoparasitic sheep tick Ixodes ricinus infestation on European Golden Plover chicks Pluvialis apricaria and implications for growth and survival. David J. T. Douglas and James W. Pearce-Higgins. Bird Study. June 2019.

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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

Ireland’s wintering waders

There’s still space for a few Knot

The island of Ireland is a great refuge for wintering waders, washed as it is by the warm waters of the Gulf Stream. It’s just a quick hop across the Atlantic from Iceland for Black-tailed Godwits, Golden Plovers, Redshanks and Oystercatchers. For birds travelling from Siberia, such as Dunlin and Grey Plovers, it’s a longer journey but one that’s well worth making.

If Ireland is such a great destination for shorebirds, why do the latest population estimates reveal a decline of nearly 20% in wader numbers in just five years?

This blog summarises the wader information, published in Estimates of waterbird numbers wintering in Ireland, 2011/12 – 2015/16 in Irish Birds. The totals in the report are split into counts for the Republic of Ireland and Northern Ireland but, given that waders don’t recognise borders, most of the comments in this blog relate to the whole of Ireland. The results for 2011-16 have been compared to the equivalent figures for 2006-11 and set in the context of the totals of wintering waders throughout the East Atlantic flyway, as combined by Wetlands International. The Irish data were collected by the amazing volunteers who make monthly, winter counts for I-WeBS (BirdWatch Ireland & National Parks & Wildlife Service) and WeBS (BTO/RSPB/JNCC in Northern Ireland).

Headline figures

Fifteen species are considered in this report. The most numerous are Lapwing and Golden Plover, which account for an estimate over 170,000 individuals between them, whilst the smallest contributions are made by Purple Sandpiper (662) and Greenshank (1317). In total, the average estimated number of waders in the winters during the period 2011-16 is 429,170 birds but it should be noted that this total excludes two widespread and common species – Woodcock and Snipe – as well as the enigmatic Jack Snipe. To update previous estimates for these three species, which were last made using distribution and abundance data collected during Bird Atlas 2007-11 fieldwork, it would be necessary to run a special inland survey. There is also some question about Lapwing and Golden Plover numbers, simply because so many of these birds are found in areas that are not covered by monthly waterbird counts.

Biggest changes

The Irish Sanderling population has increased by 13.2% in five years

The combined average winter maximum count of the 15 wader species examined in the report declined by 102,310 birds (19%) in the five-year period between 2006-11 and 2011-16. This is extremely worrying. If Lapwing and Golden Plover are excluded from consideration, as there is uncertainty about the completeness of counts, there are five species that are of particular concern; Knot numbers dropped by more than 40% and Oystercatcher, Dunlin, Redshank and Turnstone numbers by more than 20%. The Purple Sandpiper population estimate dropped by over 30% but relatively small numbers of this species are encountered around the rocky coast of Ireland. The only species to show increases were Sanderling, Bar-tailed Godwit, Black-tailed Godwit and Greenshank.

In a previous WaderTales blog, there is detailed information about population estimates for Great Britain: Do population estimates matter? In Great Britain there were similar rates of decline for Redshank and Turnstone (measured over an eight-year, rather than five-year period) but much smaller falls for Knot, Oystercatcher and Dunlin. The possible causes of the changes in Ireland are discussed in the paper in Irish Birds. They include flyway-scale declines (e.g. Knot and Curlew) and the possibility that more birds from the east are now wintering on the coasts of mainland Europe (e.g. Dunlin and Grey Plover).

European context

The table alongside gives an indication of the relative importance of Ireland, Great Britain and, together, the British Isles to the birds that use the East Atlantic flyway during the winter period. The three columns show the percentage of each species found in each of the three regions. Summarised international counts, as used in the paper, were kindly provided by Wetlands International. In the case of four species, Ireland is host to a significant proportion of the Icelandic breeding population (Oystercatcher, Golden Plover, Black-tailed Godwit and Redshank). There’s a WaderTales blog about the close link between Ireland & Iceland. Another blog – Which, wader, when & why? – summarises migration to, from and through Britain & Ireland.

Notes: As mentioned earlier, there are questions about the precision of estimates for Lapwing and Golden Plover, although the population trends are reliable. The Ringed Plover percentage seems high (98% for British Isles) but this may well reflect the fact that the Non-estuarine Waterbird Survey has uncovered significant numbers of the species on the open shores of Great Britain. These extra birds are included in the new totals for GB but not in the flyway total. The percentages for Black-tailed Godwit seem low, as discussed further down.

Ireland is particularly important for Golden Plover, Ringed Plover, Bar-tailed Godwit and Black-tailed Godwit, as well as for the Icelandic subspecies of Redshank. Greenshank is excluded because the percentages are below 1% of the flyway population for Ireland and for Great Britain.

11% of Bar-tailed Godwit on the East Atlantic Flyway spend the winter in Ireland

Although there are important populations of breeding waders in Ireland, the shores and wet fields of the island really come into their own during July and August, when the first ‘winter’ waders arrive, and they only become quiet again in April and May, when the last birds head north and east to nest. A successful breeder is likely only to be away for four or five months, meaning that these waders will spend by far the largest part of the year in Ireland. The island is even more important for immature birds. Young Oystercatchers that arrive from Iceland, Scotland or perhaps Norway when just a few months old are likely to spend the next 30 months in Ireland before making their first trip north. There is a WaderTales migration blog about the Oystercatchers that fly from Iceland: Migratory decisions for Icelandic Oystercatchers.

Curlew in the Republic

Curlew numbers in the Republic of Ireland illustrate the relative importance of the country for breeding and non-breeding populations. The winter population estimate for Curlew in the Republic is 28,300 but the most recent survey conducted by BirdWatch Ireland and NPWS, as summarised in the WaderTales blog Ireland’s Curlew Crisis, reveals that the number of breeding birds has crashed to just 138 pairs. Accounting for young Irish birds that have not started to breed, and even if we assume that all Irish birds stay in the country for the winter, then the total number of home-grown Curlew seen in non-breeding flocks is at most about 400. This means that every winter flock of 70 Curlew will contain an average of just one Irish bird. Far more deliver their curl-ew calls with a Scottish, Finnish or Swedish ‘accent’. The map below shows the migration pattern for Curlew ringed in or found in Britain & Ireland.

Black-tailed Godwit

In the table above, it looks as if 18% of Iceland’s Black-tailed Godwits spend the winter in Ireland. This is probably an underestimate of the importance of the Republic of Ireland and Northern Ireland to the species. The flyway total for Black-tailed Godwit is given as between 98,000 and 134,000 in the Irish Birds paper and the percentage figure is based on 110,000. These three figures are almost certainly too high, as they build upon country-based estimates that have subsequently been revised. The true figure is likely to be around 60,000 to 65,000 (J. Gill pers. comm.), which would suggest that the maximum winter count in Ireland of 19,800 represents at least 30% of the islandica Black-tailed Godwits. Add in extra birds that moult in Ireland in the autumn, before moving further south to countries such as Portugal, and other birds that spend spring months on the island, and Ireland becomes even more important for Black-tailed Godwits!

Most birdwatchers might associate flocks of waders with estuaries but Black-tailed Godwit is an excellent example of a species that also relies on inland fields, either close to estuaries or along river valleys. Whilst undertaking PhD research on Black-tailed Godwits in south-east Ireland, Daniel Hayhow showed that there is insufficient time to find enough estuarine food during the mid-winter tidal cycles, with birds topping up their resources on grassland. You can read more about the energetic consequences of choosing to winter in eastern England, Portugal and Ireland in this blog: Overtaking on migration. Site designation and planning decisions need to take account of the grassland feeding requirements of Black-tailed Godwits and other waders that do not spend all of their time on estuaries, particularly Curlew.

Conservation implications

Some of the issues facing waders may be related to threats that species face in the breeding grounds. However, it may be easier to introduce measures that provide better protection and feeding opportunities in the wintering area, as ways of maintaining populations through the non-breeding season, than it is to deal with problems in the High Arctic. (Although we can all help by reducing carbon emissions, in order to minimise global warming, of course).

Reading the report, I was reminded of the need to consider a range of conservation issues:

• Care needs to be taken when considering shoreline developments. These can directly remove habitat or squeeze the width of the intertidal zone.
• Increased harvesting of shellfish can affect species such as Oystercatcher and Knot and brings risks of introducing alien species and diseases.
• In the drive to cut carbon emissions, tidal, wave and wind power developments need to be sited in appropriate places.
• Off-shore harvesting of growing kelp beds has been suggested, as a way of producing fertiliser and biofuels. This process could reduce protection for beaches and change the availability of resources for species such as Turnstone and Sanderling.
• Grassland areas need to be considered (and not just estuaries) when planning protection for species such as Curlew and Black-tailed Godwit.

Paper

Estimates of waterbird numbers wintering in Ireland, 2011/12 – 2015/16. Brian Burke, Lesley J. Lewis, Niamh Fitzgerald, Teresa Frost, Graham Austin, and T. David Tierney. Irish Birds No. 41, 1-12.

There is a complementary paper in British Birds, covering Great Britain. The wader information is summarised in this blog: Do population estimates matter?

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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

Do population estimates matter?

How many waders spend the winter in Great Britain? The answer is provided within an article in British Birds entitled Population estimates of wintering waterbirds in Great Britain. It includes all the wader species from Little Stint to Curlew that are covered by the Wetland Bird Survey  This survey is based on monthly counts that take place at about 2000 wetlands and coastal sites. The main aim is to monitor the rise and falls in numbers over time.

Please note that Northern Ireland WeBS figures are included in a separate blog covering the island of Ireland that was published in Irish Birds.

Why do we need to know the total number of birds in Great Britain?

• If we count the number of Curlew and we have a figure for the European population then we know that Great Britain is responsible for nearly 20% of Europe’s Curlew each winter, thereby strengthening the case for national conservation action;
• If we have a national figure, then we know that a flock of 2000 Black-tailed Godwit represents (as it turns out) over 5% of the British total, which is a useful criterion when assessing the conservation importance of individual sites;
• Population totals help to put annual percentage changes into context;
• And simply because people ask questions such as “how many Greenshank are there in the country during the winter?”

So, here’s the bottom line. In their 2019 review of waterbird numbers in British Birds, a team from BTO, WWT, JNCC & RSPB reveal that an estimated total of 4.9 million waders spend the winter in Great Britain. For several species, GB holds a third or more of the East Atlantic Flyway population!

Making the counts

The population estimates owe a lot to those who undertake monthly Wetland Bird Survey (WeBS) counts on estuaries, lakes and waterways, during the winter months, year in and year out. Counts from the period 2012/13 to 2016/17 are used in the population estimates that form the basis for the 2019 review. WeBS data have many other uses, as you can read here: Wetland Bird Survey: working for waders.

For species of wader that also make use of the open coast, the Non-estuarine Waterbirds Survey of 2015/16 (or NEWS III) provided additional data, updating the NEWS II figures from 2006/07.

The vast majority of our wintering Purple Sandpipers are found on open beaches and rocky shores, as well as large numbers of Turnstone, Ringed Plover and Sanderling, together with significant numbers of Oystercatcher, Curlew and Redshank. There’s more about NEWS in this slightly dated blog: NEWS and Oystercatchers for Christmas.

The last assessment of winter wader populations was made by the Avian Population Estimates Panel and published in British Birds in 2013 as Population estimates of birds in Great Britain and the United Kingdom (APEP3). In here, estimates for waders were largely based on WeBS data for the period 2004-09 and NEWS II. The new assessment is presented as Population estimates of wintering waterbirds in Great Britain and also published in British Birds. It uses WeBS information for the period 2012-17 and NEWS III data. Effectively, there is an 8-year or 9-year difference between the two sets of figures.

The biggest losers

Great Britain is extremely important in the context of the East Atlantic Flyway, as is obvious from the fact that the area holds nearly five million waders. The WeBs counts already monitor the ups and downs on an annual basis but this review provides an opportunity to turn the percentages into actual numbers. It is concerning that, over a period representing less than a decade, the average maximum winter count for six of the species that were surveyed dropped by a total of over 150,000. These big losers were Knot, Oystercatcher, Redshank, Curlew, Grey Plover and Dunlin, ordered by number of birds lost, with Knot seeing the biggest absolute decline.

In preparing the new estimates for the British Birds paper, an opportunity was taken to refine the way that populations are calculated, based on Use of environmental stratification to derive non-breeding population estimates of dispersed waterbirds in Great Britain, by Verónica Méndez et al. The new methodology explains some of the differences between percentage changes reported by WeBS and the percentage changes obtained by comparing the latest population estimates to those in APEP3.

The Knot estimate dropped from 320,000 to 260,000. This decline is bigger than might be expected from the counts that take place at sites covered by WeBS, being larger than the ten-year decline of 14% reported in the last WeBS report. Knot are mobile species within the North Sea and Atlantic Coast wintering area and it is possible that British losses may be explained, at least to some extent, by redistribution.

The drop in Oystercatcher numbers from 320,000 to 290,000 appears to be less than 10%, compared to a ten-year decline of 12% on WeBS. Improved analysis of NEWS data helped to add some more birds to the open-coast estimate so the 10% fall may underestimate the seriousness of the Oystercatcher situation. The 25-year Oystercatcher decline on WeBS is 26%, which is not surprising if you look at the changes to breeding numbers in Scotland, where most British birds are to be found. There’s more about this in: From shingle beach to roof-top.

The Redshank decline of 26,000 is higher than would be predicted from WeBS figures, suggesting a drop of over 20% since APEP3, rather than ‘just’ 15% for the ten-year WeBS figure. This is a species that also features strongly in the Non-estuarine Waterbird Survey and that might explain the difference. Wintering Redshank are mostly of British and Icelandic origin, with the Breeding Bird Survey (BBS) suggesting a ten-year decline of 24% in our British breeding birds.

The Curlew is now globally recognised as near-threatened. The latest winter estimate is 120,000, down from 140,000 in APEP3. The new total represents between 14% and 19% of the European population, which means that we have a particular responsibility for this much-loved species. Only the Netherlands holds more wintering Curlew than Great Britain. Is the Curlew really nearly-threatened? is one of several blogs about Curlew in the WaderTales catalogue at www.wadertales.wordpress/about .

It has been suggested that the long-term declines of Grey Plover and Dunlin  may be associated with short-stopping, with new generations of both species wintering closer to their eastern breeding grounds than used to be the case. WeBS results indicate a 31% drop in Grey Plover and a 42% drop in Dunlin, over the last 25 years. There was a loss of 10,000 for both species between APEP3 and the new review, representing declines of 23% and 3% respectively.

The biggest winners

There are several big winners in the period between APEP3 (2004-09) and the new review (2012-16), although, in some cases, not all is as it seems.

The Avocet has seen further dramatic gains. with the estimated wintering population rising to 8,700. The increase is not quite as big as might have been expected, based on the 43% rise seen in ten years of WeBS counts, but it is still a dramatic continuation of a 40-year trend.

The numbers of Bar-tailed Godwit and Ringed Plover are both substantially higher but at least a proportion of each of these changes is linked to the better coverage and more sophisticated sampling methods that were discussed earlier. Bar-tailed Godwit increases may also reflect redistribution around the North Sea.

One of the consequences of improved statistical techniques, as used this time around, is the apparent decline in the estimated population of Black-tailed Godwit. The new figure of 39,000 is 4,000 smaller than in APEP3, despite the fact that the WeBS graph clearly shows an increase. Interpolation using WeBs figures suggests that the earlier population estimate should have been 31,000, rather than 43,000.

Sanderling from Greenland spend the non-breeding season as far south as South Africa but  increasing numbers of birds are wintering in Great Britain and Ireland (25% increase in 8 years in GB and 13% in 5 years in Ireland). Interestingly, survival rates of English birds are just as high as those in Namibia. The losers are birds that spend the non-breeding season in equatorial Africa, as you can read here; Travel advice for Sanderling.

There are other winners too, as you can read in the paper. At the start, I posed the question “how many Greenshank are there in the country during the winter?”.  The answer is 810, representing an increase of 200 since APEP3. The vast majority of these wintering Greenshank are birds from the population that breeds in northern Scotland, as you can read in Migration of Scottish Greenshank.

Game species

The estimates for the three wintering waders that are still on the UK quarry list have not changed since APEP3 (published in 2011) as there are no new data available.

Golden Plover: The winter estimate remains as 400,000, as there has been no comprehensive, winter survey since 2006/7. Large numbers of Golden Plover arrive from Scandinavia, Europe and Iceland in the late summer, joining the British birds that choose not to migrate south or west. The GB breeding population is probably less than 50,000 pairs. Most breed in Scotland which has seen a breeding decline of 5% in the period 1995 to 2018 (BBS). Golden Plover is still ‘green listed’.

Snipe (Common): The winter estimate remains as 1,100,000 – a figure that was acknowledged in APEP3 as being less reliable than that of most species. At the same time, the GB breeding population was estimated as 76,000 pairs, indicating at least a 4:1 ratio of foreign to British birds, and that does not take account of the number of British birds that migrate south and west. Snipe are ‘amber listed’ but BBS suggests a recent increase of 26% (1995-2018). There is a WaderTales blog about  Snipe and Jack Snipe.

Woodcock: The winter estimate remains as 1,400,000 – another figure that is not considered to be particularly precise, with much variation between years. The diminishing breeding population is dwarfed by winter numbers, as you can read in this WaderTales blog, with increased attention being given to ways to afford better protection of red-listed, British-breeding birds.

Many of the Golden Plover, Snipe and Woodcock that spend winter in Great Britain are birds that breed in Fennoscandia (Finland, Sweden & Norway). The latest assessment of breeding numbers shows that populations of all three species are stable. See Fennoscandian Wader Factory.

January counts

The paper in British Birds also includes a table of January population estimates, to provide data that are comparable to mid-winter counts in other countries. These figures are used in waterbird monitoring for the International Waterbird Census for the African Eurasian Flyway. The main table (and figures mentioned above) are average maximum winter counts (in the period September to March). Black-tailed Godwit is one species that illustrates the difference, with a mean of 30,000 in January and a mean peak count of 39,000. Having moulted in Great Britain, some Black-tailed Godwits move south to France and Portugal in late autumn, returning as early as February. January counts are therefore substantially lower than early-winter and late-winter counts. There is more about the migratory strategy employed by Black-tailed Godwits that winter in southern Europe in Overtaking on Migration.

Looking forward

The authors have done a tremendous job. They have refined the way that estimates are calculated, they have combined the results from WeBS and NEWS III, and they have delivered population estimates for 25 wader species and many more other species of waterbirds. These population estimates will be used in conservation decision-making until the next set of numbers becomes available. Meanwhile, thousands of birdwatchers will count the birds on their WeBS patches in each winter month, every year. Without them, this paper could not have been written.

Before the next assessment, there will need to be another NEWS survey, to check up on species that use rocky and sandy shore birds, such as Purple Sandpipers, Turnstone and Curlew. Hopefully, there will also be a dedicated survey to assess Lapwing and Golden Plover numbers and perhaps we might find a way to refine the old estimates for Woodcock, Snipe and Jack Snipe.

Paper

Population estimates of wintering waterbirds in Great Britain. Teresa Frost, Graham Austin, Richard Hearn, Stephen McAvoy, Anna Robinson, David Stroud, Ian Woodward and Simon Wotton. Published in British Birds Volume 112. March 2019.

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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

Designing wader landscapes

Much has been written about the negative impacts of agriculture on breeding birds – but farming can be good for some species. In Iceland, where high-input agriculture is relatively recent, breeding waders are commonly found in nutrient-rich environments that are associated with increased production. How can high breeding densities of waders be maintained, as farming continues to expand and intensification increases?

In her paper in Agriculture, Ecosystems & Environment Lilja Jóhannesdóttir investigates the distribution of breeding waders across landscapes with varying amounts of highly-cultivated fields and semi-natural areas. She discovers that, in some circumstances and at an appropriate level, adding cultivated land within a broader mosaic of habitats may benefit breeding waders. Is this a model system that provides clues as to how to design landscapes that can support sustainable breeding wader populations in other parts of the world?

The waders of Iceland

Breeding populations of waders in Iceland (AEWA report)

Iceland is a hot-spot for breeding waders, holding half or more of Europe’s Dunlin, Golden Plover and Whimbrel, in a country that is a bit smaller then England. The paper at the heart of this blog is written by Lilja Jóhannesdóttir, who worked with colleagues from the University of Iceland, the Agricultural University of Iceland, the University of East Anglia (UK) and the University of Aveiro (Portugal). They investigated how different ways of increasing agricultural productivity might impact upon these species, and others such as Black-tailed Godwit, Redshank and Snipe.

Much of Iceland’s upland interior is not suitable as farmland but there is still plenty of room for agricultural expansion. Only 7% of the area between sea level and an elevation of 200 m is currently under cultivation but it is estimated that it would be possible to increase this to 63% – an eight-fold extensification. Icelandic lowlands currently comprise a fine-scale mosaic of open semi-natural habitats and cultivated fields (primarily for silage production to feed animals), making most of the landscape much more heterogeneous than in countries with a longer history of commercial farming.

Two previous WaderTales blogs have already shown that:

• Most Icelandic farmers expect to increase the amount of land that they cultivate but may be prepared to modify their plans in ways that can accommodate waders – which they like seeing on their estates. Do Iceland’s Farmers care about wader conservation? is based upon a paper published in Ecology and Society.
• Generally, there are lower densities of breeding waders in the most highly-managed fields but with some exceptions. For instance, in the west of the country, where underlying soil productivity is lower, waders are more common in intensively managed fields later in the season, when chicks are more mobile. Farming for waders in Iceland is based upon a paper in Animal Conservation.

Given that farm production is predicted to increase, that farmers like breeding waders and that some intensively-managed fields can be attractive to waders, is it possible to design farmed landscapes that will work for birds and farmers?

Increasing inputs and reducing heterogeneity

Globally, the expansion and intensification of agriculture has altered landscapes and the associated homogenisation has greatly influenced bird abundance and reduced biodiversity. Populations of numerous species, particularly specialist species, have declined, as agriculture has expanded, while generalist species have often thrived in agricultural habitats.

There is no shortage of examples in which highly intensively managed farmland is shown to be bad for breeding waders. In the monoculture hay-meadows of the Netherlands, Black-tailed Godwit productivity is really low, for instance. These fields have been drained, fertilised and re-sown, in order to create easily-managed carpets of single-species grass that can be cut several times a year. There is more about this in this paper by Roos Kentie.

Although there are some areas of Iceland in which farming is quite intensive, there are many others where farmers have a lighter touch. For instance, nutrient-poor dwarf birch marshes may occasionally be grazed by sheep in the summer but these areas have never received applications of artificial fertiliser. At this end of the intensification continuum, increasing agricultural operations may have benefits for breeding waders. When a patch of rough grazing is ploughed and turned into a hay meadow, the addition of fertilisers can potentially increase soil fertility and create an attractive place for waders to feed. A hay meadow within a local area that is dominated by dwarf birch marsh could effectively increase the heterogeneity (& nutrient-richness via spill-over) of the local area, albeit in an artificial way. In the UK, Golden Plovers breeding on moorland are known to travel up to 7 km to feed on fertilised hayfields with high earthworm densities. This paper by James Pearce-Higgins & Derek Yalden in IBIS provides a nice example of how low intensity agriculture can provide resources for waders in the wider landscape.

Researching waders and landscapes

Lilja’s work in the Southern Lowlands of Iceland focused upon understanding how agriculture influences breeding wader densities and how these relationships might influence future change. At its heart were counts of adult waders encountered along 200 transects (totalling over 100 kilometres) within semi-natural habitats, visited at several stages during the breeding seasons of 2011 and 2012.

As well as counting birds, Lilja categorised habitats within 500, 1000, 1500 and 2500 metres of the transects, which she called buffer area in the paper. Interestingly, and usefully for later analyses, the distribution of different habitat types is pretty uniform across these scales, in this part of Iceland, with little substantial difference according to elevation. In the diagram below, the 200 transects (a) have been split between those below 50 m above sea level (b) and those higher than 50 m (c).

Landscape-scale effects

To fulfill the various demands of parents and their offspring, waders need diverse resources on or near their territory. An adult can feed a kilometre or more away from its nest, between incubation bouts, and chicks are mobile from an early age. Tagging has shown that young Black-tailed Godwits can move up to 3 km in the first five days of life, just to give one example. In this open landscape, breeding success is likely to be a function of habitat availability at a broad scale. This is explored in a WaderTales blog about nesting Whimbrel.

Using data collected from these 200 lowland transects, Lilja was able to establish relationships between breeding wader densities and the amount of cultivated land and wetland in the surrounding landscape. These two habitat types were considered because future agricultural expansion is likely to take place on drained wetlands that have high conservation value. In her analyses she assessed the extent to which the amount of cultivated land in the surrounding landscape affects wader densities on semi-natural land, and then considered the potential effects of future agricultural expansion on wader populations. There was substantial variation in the density of all of the six most common wader species recorded on the transects, ranging from 0 to 284 birds/km².

Lilja found that wader densities in semi-natural habitats were consistently greater when the surrounding landscapes had more wetland, at scales ranging from 500 m to 2500 m, indicating the importance of wetland availability in the local neighbourhood. However, the effects of cultivated land in the surrounding landscape varied with fertility and landscape structure, which was largely defined by altitude.

• In fertile, low-lying coastal areas (from sea-level to 100 m altitude), wader numbers declined with increasing amounts of cultivated land (and the lowest densities occurred in areas dominated by cultivation). This suggests that further conversion of semi-natural habitats into farmland is likely to severely impact waders in low-lying areas.
• In less fertile habitats at higher altitudes (between 100 m and 200 m), the lowest densities occurred in areas without cultivated land. This suggests that additional resources provided by cultivated land may have a more positive affect in the less-fertile, higher altitude areas.

The relationships between the areas of wetland and agriculture in the surrounding landscape and the density of waders vary between species, as you can read in some detail in the paper. A few highlights are:

• With increasing area of cultivated land, densities of Golden Plover, Dunlin and Whimbrel declined significantly at lower altitudes but increased at higher altitudes. These are the three species that would appear to respond most positively to the addition of pockets of cultivated land within a semi-natural matrix of less fertile land, that tends to be found at higher elevations.
• Higher amounts of wetland were associated with increased densities of Dunlin and Black-tailed Godwit, but lower densities of Redshank. Golden Plover numbers were unaffected by amount of wetland in the surrounding landscape.
• Whimbrel densities increased with wetland area, at higher altitudes. Wet patches have been shown to be very important to Whimbrel chicks, as you can read in this WaderTales blog about research in Shetland.
• At lower altitudes, Snipe densities increased with the amount of wetland area in the local vicinity. This relationship was less pronounced at higher altitudes, which tend to be less effectively drained and hence generally wetter.

What now?

Changes in Icelandic landscapes are to be expected in the coming years, as most farmers intend to increase their areas of cultivated land. This expansion will inevitably have impacts upon the internationally important breeding wader populations of Iceland but the level of such impact will depend on where the expansion will occur. This paper shows that increases in the area of cultivated land at lower altitudes in Southern Iceland are more likely to result in declines in wader density than in less fertile areas, which are usually found at slightly higher altitudes (still under 200 m above sea level). An important next step will be to identify the landscape structures and scales of management that can continue to support high densities of breeding waders.

Given the international importance of Iceland as a home for breeding waders it would be nice to think that this paper can be used to develop national land management policies that can prevent the unintended loss of species such as Golden Plover and Snipe, which landowners value and wish to preserve. At the farm and community level, the paper highlights the key importance of maintaining the complex and heterogeneous landscapes of lowland Iceland, retaining as many as possible of the remaining wetland patches and pockets of semi-natural land within even the most intensive of farming areas.

The paper may well be of interest to conservationists who are struggling to reverse wader declines in other parts of the world. In Southern Iceland, where 7% of the land is being farmed relatively intensively within a fine scale mosaic of both wet and dry semi-natural habitats, it is possible to support hundreds of waders per square km across the wider countryside. Can this situation be replicated across large tracts of land in other countries?

Take home message and paper

This paper provides a useful reminder that the links between land use changes and biodiversity implications can be highly context-dependent. Further agricultural conversion of wetlands and rough grazing areas in the fertile low-lying areas of Iceland is likely to be detrimental for breeding waders, but such effects may be less apparent in less fertile, higher altitude areas. Here, the conversion of some land from rough-grazing to hay meadows may provide feeding opportunities off-territory for Dunlin, Golden Plover and Whimbrel. The scale at which the addition of cultivated areas is beneficial to breeding waders has yet to be determined.

This paper is published as:

Interacting effects of agriculture and landscape on breeding wader populations. Lilja Jóhannesdóttir, Jennifer A. Gill, José A. Alves, Sigmundur H. Brink, Ólafur Arnalds, Verónica Méndez and Tómas Grétar Gunnarsson https://doi.org/10.1016/j.agee.2018.11.024

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Graham (@grahamfappleton) has studied waders for over 40 years and is currently involved in wader research in the UK and in Iceland.  He was Director of Communications at The British Trust for Ornithology until 2013 and is now a freelance writer and broadcaster.

Starting moult early

In waders (shorebirds), the main moult (molt) usually takes place after the migration that follows the breeding season. Golden Plovers adopt a different strategy, starting wing moult while still nesting. Given that these adult birds are not going to fly anywhere any time soon, this seems like a very efficient strategy. So, why do Icelandic and Scandinavian Golden Plovers moult differently? Is this a reflection of available resources?

The post-breeding moult

Moult is an energetic process, especially the post-breeding moult, which includes a change of all of the wing and tail feathers. To complete the whole process, birds ideally need to find a three-month period when resources are good, climatic conditions are benign and there is no need to migrate. For birds on the East Atlantic Flyway that spend the non-breeding season in Europe, moult typically takes place after the breeding season and before days get shorter and the weather gets colder.

One place with plenty of food-rich mud is the Wash, in eastern England. Here, up to 300,000 waders gather each autumn, including Knot from Greenland and Canada, Grey Plover from Siberia and Curlew from countries such as Finland. A relatively small proportion are juveniles, which will only moult their body feathers, but there are probably at least 200,000 waders in full moult at this time of year – dropping and growing a grand total of perhaps a billion feathers between them. Some populations use the Wash as a feeding station, before moving on to moult in their wintering grounds, but this is a minority. This group includes taymyrensis Bar-tailed Godwits (more about these birds here) and schinzii Dunlin, which will travel further south, to Africa.

Golden Plovers start their moult during or before the incubation period

There are Golden Plovers spread across the autumn mud-flats of the Wash too, made up of a mixture of birds that have bred in Britain, Europe, Scandinavia and Iceland. Although they end up in the same winter flocks, their moult strategies are different. Recent research by Paula Machín and colleagues has focused upon how breeding season conditions impact upon the moult strategy of two distinct Golden Plover populations, birds breeding in Scandinavia and Northern Russia and others breeding in Iceland. The resulting paper is published in the Journal of Avian Biology.

Conditions at the breeding grounds and migration strategy shape different moult patterns of two populations of Eurasian golden plover Pluvialis apricaria Paula Machín, Magdalena Remisiewicz, Juan Fernández-Elipe, Joop Jukema & Raymond H.G. Klaassen

Icelandic Golden Plovers

Up to one million Golden Plovers arrive in Iceland each spring, mainly from Ireland and western parts of the United Kingdom. This is estimated to be nearly half of the European breeding population. Iceland might seem small, when compared to the vast land-mass of the European continent, but it’s a haven for waders. This status is threatened by the spread and intensification of lowland farming, increased afforestation and by the ‘summer cottage’ industry – but those are stories for another day.

The Heiðlóa (Golden Plover) is a welcome sight and sound at the end of the Icelandic winter. The first migrants appear about 23 March and nesting can commence as early as 26 April. The usual clutch size is four eggs, with both parents sharing incubation duties. Some first nests are lost, due to predation, but females can lay another clutch. Joop Jukema studied Golden Plovers nesting near Selfoss in the Southern Lowlands of Iceland, timing his captures of nesting birds to coincide with the later part of the incubation period. He was able to assess the progress of moult by scoring the growth of the primary feathers and to work out when each bird would have dropped its first primary. The estimated mean start date of primary moult for males was 19 May (95% confidence interval 27 April – 10 June) with females starting an average of 9 days later, on 28 May (95% confidence interval 6 May – 19 June). On average, males started to moult their primary feathers nine days before the start of incubation, while females started to moult at the same time as incubation began. Potentially, hormone changes associated with the stage of the breeding season could be linked to the onset of moult.

Icelandic Golden Plovers complete their moult prior to departure from the country. By making catches of birds in late August and early September, it was possible to show that the primary moult period is about 100 days. No birds were caught in suspended moult, strongly suggesting that Icelandic Golden Plovers do not attempt to cross the Atlantic before they have attained a complete, fresh set of feathers.

Swedish Golden Plovers

Paula Machín’s main study site was in Ammarnäs in Sweden, on roughly the same latitude as Selfoss and hence with the same amount of daylight. Ammarnäs is colder in spring than Selfoss, not benefiting from the warmer waters of the Gulf Stream which wash the shores of southern Iceland. The average start of incubation in Ammarnäs was eight days later than in Selfoss with the commencement of moult being seventeen days later. Males started primary moult three days after the start of incubation, with females starting twelve days into incubation. It is interesting to note that the difference in timing of the two sexes is nine days, just as in Iceland.

Female Golden Plovers left the Ammarnäs breeding territories at the end of July. From observations of females caught on their nests, it seemed likely that individual females were not starting the moult of their outer primaries, typically completing the moult of primary four and not dropping primary five. Males stayed with chicks for an extra fortnight. Given the longer period of time available to males, it is likely that they were able to moult more primary feathers than their partners, prior to departure from the area.

Catching birds during the chick-guarding phase or just before migration is very difficult but Raymond Klasssen and colleagues were able to study birds on similar strategies by catching birds at Lund, in southern Sweden, and in the Netherlands. Here, birds from Swedish breeding areas and further afield gather to moult, recommencing primary moult at the point at which it had been suspended. Inspecting moulting birds in the period August through to November enabled the research team to monitor the second part of the moult of Continental birds. Given that the distance between breeding areas and these staging areas is relatively short and that tracking showed that it could be completed in one or two days, it is possible that adults might be able to migrate while in active moult.

Spot the difference

Overlapping the breeding and moulting period is rare in migratory birds but it makes sense in a time-constrained annual cycle. The research team suggest that Icelandic plovers presumably need to initiate moult early in the season, in order to be able to complete it at the breeding grounds. This is not an option for Continental plovers, as their breeding season is much shorter, due to a harsher climate and an earlier drop-off in the number of arthropods, their main food source. These Golden Plovers cannot delay the start of the moult period until after the autumn migration because there is insufficient time to compete a full moult in areas such as Lund or the Netherlands, prior to the onset of winter frosts. The fact that Golden Plover are largely associated with farmland, rather than estuarine sites, may make them more susceptible to sub-zero temperatures than, for instance Grey Plover.

When incubating and looking after chicks, Icelandic and Swedish Golden Plovers were able to moult at the same rate. However, there were differences in the second part of the primary moult season. Away from their territories, Icelandic birds continued to moult at the same rate as previously but, having moved to Lund or the Netherlands, Continental birds could moult twice as fast as before. The availability of earthworms in these staging areas may make it easier to acquire resources for the energetically-expensive moult process.

Despite the faster rate of moult of Continental birds in the later period of their moult, the total period of primary moult is longer than that of Icelandic birds. Birds completing their moult in Iceland took an average of 100 days to replace their primaries, whilst Swedish-breeding birds took 16 days longer. This difference may be associated with the time taken to complete the first stage of moult, prior to the migratory flight away from the breeding sites.

A key finding of this paper is that splitting the moult period extends the total period of primary moult. For Swedish breeders, this is the best option, however, as there is insufficient time to complete autumn moult close to the breeding grounds or after the breeding season. As the authors conclude, “Meeting the energy demands of breeding, moult, and migration calls for different timing and spacing of these events in their annual cycle, adjusted to conditions at their breeding and stopover sites, and to their migration strategy.”

There’s more from Paula Machín and her colleagues on this blog-site: https://overthetreeline.wordpress.com/

More moult

There are more WaderTales blogs about moult: 

• The not-so Grey Plover includes general information about wader moult and talks about some of the stresses that may occur.
• Lapwings can moult while on migration. Read more in Lapwing Moult, which also talks about how to study moult without catching birds.
• Bar-tailed Godwit; migration and survival mentions the different strategies of two subspecies of Bar-tailed Godwit, both of which can be found on the Wash in autumn.

I look forward to future papers about moult strategies to add to WaderTales. Here’s a list of the 60+ blogs that are already available: https://wadertales.wordpress.com/about/

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Graham (@grahamfappleton) has studied waders for over 40 years and is currently involved in wader research in the UK and in Iceland.  He was Director of Communications at The British Trust for Ornithology until 2013 and is now a freelance writer and broadcaster.