Conservation beyond boundaries

When an environmental impact assessment concludes that only a small number of shorebirds will be affected by a new airport, because relatively small flocks are counted during field surveys, is there an assumption that the birds encountered are always the same individuals? What if different shorebirds use the same patch of mud at different stages of the tide, at different times of day or in different seasons? How many birds might really be affected?

In a 2023 paper in Animal Conservation, Josh Nightingale and colleagues investigate the movements of colour-marked Black-tailed Godwits, to see how much they fly into, out of and around a Special Protection Area, to work out how well an Environmental Impact Assessments (EIA) might assess the importance of a site that has been scheduled for development. They call for more use to be made of movement data, to assess the total impact of new developments proposed for estuaries and elsewhere.

Impact assessments

The pressures on estuaries have never been greater, as humans turn to them for transport, food, energy and to create land for new developments, such as airports. These muddy havens might be protected by national and international statutes but legally-enforceable conservation designations tend to melt away when there is money to be made, as discussed in Tagus Estuary: for birds or planes?

Developers often argue that they only want to use a small area; taking a bite out of an estuary may seem to affect only 10% (for instance) of a Protected Area – and leave 90%. Counting the birds that use that particular section might show whether this ‘10%’ contains more or fewer birds than expected and indicate whether an unacceptably high proportion of the species for which the Protected Area was designated might be affected. How much more can be learnt if we consider movements that take place within an estuary, the waves of birds that use the estuary in different seasons, the importance of the site within a Flyway and linkages to breeding areas?

Newly-ringed Black-tailed Godwit

Thanks to ringing and colour-ringing, we know that Protected Areas can form networks of sites used by individuals. For example, an islandica Black-tailed Godwit might moult on the Wash (Eastern England), spend the winter on the Tagus (Portugal) and spring in Morecambe Bay (Northwest England). Thousands of limosa Black-tailed Godwits that winter in West Africa spend two months in the Tagus in spring, where they will be seen alongside Grey Plovers from Siberia, Turnstones from Canada and Bar-tailed Godwits from the Arctic regions of Scandinavia. Our knowledge of the strength and complexities of these networks is deepening, still further, thanks to satellite tracking, and the same technology provides the potential to understand more about within-estuary movements too.

Network Analysis

Josh Nightingale measures a Black-tailed Godwit with the Tagus Estuary Ringing Group

Information on movements of individual birds, generated by ringing, together with the development of methods such as network analysis, provides a framework in which researchers can assess the importance of Protected Areas – or threatened parts of Protected Areas. Network analysis is the study of connections. In this context, a network is simply a collection of sites, dubbed nodes, linked by connections known as edges, which may have varying directions or strengths.

In pictorial terms, ‘thicker’ edges indicate that more birds connect two nodes. When assessing connectivity, the researchers considered the number of other nodes each node is linked to and the variety of pathways between nodes.

Despite being powerful and flexible, network analysis is currently used less by conservation practitioners than by academics, and much of their work is at landscape- or regional-scales. In a 2023 paper, Josh Nightingale and his colleagues adjust the focus, to see how network analysis might explain what is happening within and around a Protected Area – information that is valuable when trying to assess conservation implications of planning decisions. They use movement data to reveal the range of sites used by individuals, and thus the susceptibility of those individuals to a local development.

Network analyses can reveal which sites are most important to population-level connectivity, or the impact on connectivity of losing one or more sites. These results can be combined, to calculate the ‘impact footprint’: how many individuals use the impacted area and how do their movements connect with neighbouring sites (inside or outside the Protected Area). By representing movements between sites in a network, practitioners can gain a more accurate picture of how the effects of a localized impact may be felt at connected sites. This information can then be used to determine whether populations or habitats are impacted, and thus inform policy decisions.

Applying a Network Analysis framework to the Tagus

Black-tailed Godwits feeding along the shoreline of a built-up area in the Tagus estuary

The Tagus Estuary, on whose banks Lisbon sits, is Portugal’s largest wetland and the country’s most important site for many waterbird species. Part of the estuary is designated under the EU Birds Directive as a Special Protection Area (SPA), with a smaller Ramsar Site at its core. The SPA excludes several of the estuary’s high-tide roosts, which consequently lack legal protection and are vulnerable to development, erosion and other threats.

The Portuguese Environment Agency has issued an Environmental Licence, approving plans to construct an international airport in the heart of the Tagus estuary, on a site overlapping part of the SPA. The Environmental Impact Assessment (EIA), conducted for the development, considers the main threat to bird conservation to be noise disturbance, with an aeroplane taking off or landing every 2.5 minutes and flying at low altitude (<200 m) over the SPA (see article by José Alves in Wader Study). Such disturbance can cause birds to take flight, with consequent increases in daily energy expenditure. The effect can be seen even when the airport has been operational for decades (van der Kolk et al). Airport-related disturbance close to wetlands has been shown to influence the distribution of waterbird communities (Celdrán & Aymerich).

The predicted impact of the Montijo airport development varies, depending upon the threshold of noise sensitivity assumed. Although it has been shown that waterbirds alter their behaviour when subjected to noises above 50 dB(A) (see Wright et al., the study used as a reference for the EIA), the Environmental Licence for Montijo Airport only considers relevant areas impacted by over 65 dB(A). Crucially, movements of birds within, to or from the Protected Area were not considered.

To assess how network analysis might explain the real impact of the airport, Josh Nightingale and his colleagues focused on the Black-tailed Godwit, a species for which the Tagus SPA had been designated and for which a large amount of colour-ring data was available. They wanted to answer these questions:

  • How much of the local godwit population is protected by the Protected Area during the year, and how much does this overlap with the area to be impacted by development?
  • How much of the Tagus is used by birds from the Protected Area, and by birds from the area that would be affected by low-flying aircraft?
  • Which sites are linked most closely to others and how might the predicted development weaken connections between sites across the Tagus estuary?

Assembling the data sets

Black-tailed Godwit ringed as an adult in Iceland

Islandica Black-tailed Godwits have been intensively studied since 1993, when the first birds were colour-ringed. The datasets used in these analyses comprise sightings of birds ringed in Iceland, on the Wash (Eastern England) and in the Tagus (Portugal). Sightings of these colour-ringed birds have been reported by thousands of observers across the whole range of the subspecies. To get a flavour of the importance of these reports (and the dedication of birdwatchers) read Godwits and Godwiteers.

Within the Tagus Estuary, the focus was on marked birds reported between 1st January 2000 and 1st July 2020. Observations were recorded at 30 sites that were visited at least once during early and late winter, and more frequently since October 2006. To reduce the potential risk of observation error, only sites at which an individual was recorded at least twice during its lifetime were assigned to that individual.

To assess site-level noise-exposure that birds might experience, Josh Nightingale and colleagues analysed their data based on the areas in which godwits might encounter noise levels of 55 dB(A) and 65 dB(A). Full methods used in the network analysis are given in the paper.

Impacts on Black-tailed Godwits

Individual Black-tailed Godwit are conservative in their habits. At a Flyway scale, they typically use only four or five different sites per year, with this annual schedule remaining constant over up to twenty years (as discussed in Generational Change). Josh Nightingale found similar consistencies within the Tagus Estuary. Most of the 693 individually-tracked godwits were found at six or fewer sites (of the thirty available), even when observed for a decade or more. 82.8% of individual godwits used sites inside the SPA and 67.7% used sites outside the SPA. There was seasonal variation in the protection footprint of the SPA, which was three times as important in October-December as it was in January-March

In the early part of the winter, islandica Black-tailed Godwits are most likely to be found feeding on the estuary itself, with many moving to inland feeding sites, particularly rice fields, in January to March. Much of the area that would be impacted by the noise from aircraft is over the estuary so it is not surprising that the impact footprint is greater in October to December, when disturbance is estimated to affect 68.3% of the godwits, if a threshold of 55 DB(A) is used, or 40.7% based on 65 dB(A).

Godwits on the intertidal flats of the Tagus estuary

When the researchers considered individual movements within the same winter, they found that birds from within the Protected Area visited 14 out of 17 of the sites that lie outside the Protected Area. A flock of roosting or feeding birds seen well outside the SPA is very likely to include birds that also feed in the SPA. There were similar levels of connectivity from unimpacted sites across the estuary to the 55 dB(A) impact zone. The most centrally connected site, the Giganta rice fields, is outside the SPA but within the 55 dB(A) affected area – hence unprotected and highly impacted.

Take off! Part of a flock of thousands of Black-tailed Godwits taking off from a rice field in the Tagus estuary

Turning the focus to the area of the estuary that will be most impacted by aircraft noise, the researchers found that 68% of islandica Black-tailed Godwits in the Tagus will be affected. This is higher than an estimate of between 0.5% to 5.5% that was quoted in the EIA. The lower estimate is based on a 65 dB(A) threshold, rather than 55 dB(A), uses old counts and does not consider bird movements. This implies that the effect of the airport on Black-tailed Godwits will be twenty times as large as considered in planning decisions. And this discrepancy is just for one of the species for which the Tagus is designated as an SPA and Ramsar Site.

The paper contains a detailed analysis of how the protected network would be degraded by the new airport, through loss of key sites and by taking out edges that link current nodes that will be affected by disturbance. Overlaying the 55 dB(A) on the current Tagus network has the potential to reduce connectivity between sites by nearly 30%, effectively partially blocking the free flow of birds around the estuary.

Conservation implications

The footprint approach developed by Josh Nightingale is really neat. For the Tagus, it shows that over half of the colour-marked Black-tailed Godwits use sites outside the Special Protection Area, as currently defined, and that the majority of the most important sites that provide connectivity across the estuary are also unprotected.

Using Black-tailed Godwits as an example, the research team show that the EIA may have underestimated the impact of the airport by a factor of about 20. When disturbed by aircraft or scared off to reduce strike risk, more energy will be expended and some birds may permanently desert key feeding or roosting areas. Research elsewhere has shown that displacements can have temporary and even long-term effects on the survival rates of affected individuals.

To quote from the paper, “Protected Areas are a critically important conservation tool to protect populations, especially as ranges shift in response to climate change. To secure the integrity of PAs and the populations they support, we need to be able to accurately assess the impacts of developments inside and outside PA boundaries. Animal-tracking data offer exciting and feasible opportunities to assess PAs’ contributions to protecting populations of mobile species and the potential for adverse effects of external developments on PA integrity.

Conservation beyond Boundaries: Using animal movement networks in Protected Area assessment. Josh Nightingale, Jennifer A. Gill, Böðvar Þórisson, Peter M. Potts, Tómas G. Gunnarsson & José A. Alves. Animal Conservation. Doi: 10.1111/acv.12868.

Black-tailed Godwits moving to roost on the saltmarsh, as the tide covers the mud

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.

Why count shorebirds? A tale from Portugal

The Sado Estuary is one of Portugal’s most important wetlands – a key link in the chain of sites connecting Africa and the Arctic, on the East Atlantic Flyway. In a paper in Waterbirds, João Belo and colleagues analyse changes in numbers of waders wintering in this estuary over the period 2010 to 2019, with a focus on roost sites. These results are interpreted in regional and flyway contexts. The team find serious declines in numbers of Avocet, Dunlin and Ringed Plover.

Lost roost sites

The Sado Estuary became a nature reserve in 1980 and has since been classified as a Ramsar Site, a Special Protection Area (SPA – Natura 2000) and an Important Bird and Biodiversity Area. The Estuary lies about 30 km south of the much larger and more famous Tagus (or Tejo) Estuary, which is threatened by a new international airport (see Tagus Estuary: for birds or planes?). A suggestion that the Sado could provide mitigation habitat for damage done to the Tagus prompted scientists from the Universities of Aveiro and Lisbon to review the current state of the estuary.

João Belo and his colleagues used monthly data from a programme of wader surveys, conducted largely by volunteer birdwatchers. These took place between January 2010 and December 2019. Roosting birds were counted at high tide along the northern shores of the Sado estuary and any habitat changes were noted.

During the ten-year survey period, 21% of the available high-tide roost area was lost. These changes were associated with the commercial  abandonment of saltpans (with consequent increases in vegetation) and the conversion of others for fish farming (often with netting, to keep out fish-eating birds).

Results of the survey

In their paper, João Belo and colleagues focused on total numbers of waders and counts of the six most commonly-encountered species: Avocet, Dunlin, Black-tailed Godwit, Redshank, Ringed Plover and Grey Plover. They compared winter (Dec, Jan and Feb) counts in 2019 with those from 2010. This is when peak numbers of Avocet, Redshank and Grey Plover occur. Higher counts of Dunlin are made in spring, as schinzii birds returned from Africa, with peak counts of Black-tailed Godwit and Ringed Plover occurring in autumn.

The key findings are:

Ringed Plover numbers dropped by 23%
  • There was a strong decrease in the overall number of waders wintering in the Sado Estuary. This trend is mostly driven by steep declines in three of the six most abundant species: Avocet, Dunlin and Ringed Plover.
  • Avocet numbers were 42% lower in 2019 than they had been in 2010.
  • Dunlin numbers dropped by over half, with 2019 counts being only 47% of those in 2010. These are mostly dunlin of the alpina subspeciesthat breed between Northern Scandinavia and Siberia.
  • Ringed Plover numbers dropped by 23% between 2010 and 2019.
  • Redshank increased significantly between 2010 and 2019, while the population of Grey Plover was relatively stable, and it was not possible to derive a population trend for Black-tailed Godwit.

It is interesting to look at these patterns alongside data collected in Britain & Ireland, over the same period. As discussed in Do population estimates matter? and Ireland’s wintering waders, there have been major changes in wader numbers, with most species currently in decline.

At the same time that Avocet numbers have dropped on the Sado Estuary they have rocketed in the UK (here seen on the Humber Estuary)
  • Winter Avocet numbers have increased massively in Britain & Ireland. It is possible that young birds are more easily able to settle in these northern areas, now that winter temperatures are generally warmer. Declines in Portugal may reflect a northwards shift of the winter population, driven by new generations of birds.
  • Numbers of Ringed Plovers in Britain & Ireland did not change over the period 2010 to 2019 but had dropped a lot in the preceding twenty years.
  • For Dunlin, the size of the declines in Britain & Ireland are consistent with those on the Sado. It has been suggested that more young birds might be settling in areas such as the Wadden Sea, closer to Siberian breeding areas, something that may have become more possible given the reduced intensity and occurrence of freezing conditions along the east coast of the North Sea.

Regional and Flyway patterns

As discussed in the blog Interpreting changing wader counts, based on research led by Verónica Méndez, local changes in numbers are usually reflective of broader changes in population levels. Individual waders are unlikely to seek alternative wintering sites unless habitat is removed, so birds do not re-assort themselves into the ‘best’ areas when population levels decrease. Instead, there is general thinning out across all sites as populations decline. In this context, it is unsurprising that the trends in the Sado Estuary are similar to those found elsewhere in Portugal and in other Western European wintering areas.

Looking forwards

The survey data collected between 2010 and 2019 form a useful backdrop against which to monitor what might happen when (or perhaps if) a new international airport is constructed within the nearby Tagus Estuary. If some birds are displaced to the Sado, increases in numbers might be expected.

Displacement is not cost-free, as has been shown in a well-studied population of Redshanks on the Severn Estuary in Wales. When Cardiff Bay was permanently flooded, as part of a major redevelopment, colour-marked Redshank dispersed to sites up to 19 km away. Adult birds that moved to new sites had difficulty maintaining body condition in the first winter following the closure of Cardiff Bay, unlike the Redshank that were already living in these sites. Their survival rates in subsequent winters continued to be lower than for ‘local’ birds, indicating longer-term effects than might have been predicted.

These three papers are essential reading for anyone interested in the consequences of displacements caused by development projects.

Given that the Sado has multiple conservation designations, including as a Ramsar site, and that this study has shown a clear loss of available roosting areas, perhaps it is time to identify a high-tide refuge that can be fully protected and managed in ways which create a range of suitable habitats for use by long-legged and short-legged waders. A nature reserve such as this has a potential to attract birdwatchers too, with prospective increased income from tourism.

Sado International

Curlews don’t get a mention in this paper but the The Sado provides a neat link to a 2022 blog, A Norfolk Curlew’s Summer. This tale focuses on ‘Bowie’, a male Curlew that breeds in Breckland (Eastern England) and has been tracked to The Sado Estuary. In the blog, Bowie’s story stops in The Tagus but he subsequently headed further south to The Sado, where he spent the winter. At the time of writing (13 Feb 2023) he is still there but hopefully he will heading north soon.

The Sado is not only important in the winter, of course. As mentioned earlier, it is a spring stop-over for birds such as schinzii Dunlin, heading north from Africa to Siberia, and a moulting/staging site for waders heading south in late summer. Tracking and colour-ringing are telling more of these stories, with links to countries as far north as Canada and Siberia and as far south as South Africa.

The overgrown embankments within the former saltpans no longer provide suitable roosting sites for waders

Keep counting

The Sado story could not have been written without the work of volunteer counters who collect monthly data during the winter months, on the Sado Estuary, across Portugal and on the wider East Atlantic Flyway. These monitoring efforts are essential when attempting to track changes in wader populations, especially when extra information can indicate links to habitat changes, as is the case in the Sado. The international picture is painted using Flyway information generated using January counts that are developed by the Institute for Nature Conservation and Forests (ICNF).

Here is a link to the paper:

Synchronous declines of wintering waders and high-tide roost area in a temperate estuary: results of a 10-year monitoring programme. João R. Belo, Maria P. Dias, João Jara, Amélia Almeida, Frederico Morais, Carlos Silva, Joaquim Valadeiro & José A. Alves. Waterbirds. doi.org/10.1675/063.045.0204

Birdwatchers that volunteered to survey roost-sites gather for a team photo

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.

Curlew nest survival

The Eurasian Curlew is designated as ‘Near-Threatened’ by IUCN/BirdLife. It is Red-listed in the UK,  largely due to a rapid decline in breeding numbers. In this context, the fact that there are a few pink squares (indicating increased numbers) on the map showing breeding abundance change between 1988-91 and 2008-11 looks encouraging.

Why are Curlew doing relatively well in some areas and might these conditions be replicated? In a 2023 paper in IBIS, Harry Ewing examines how nest survival varies within Breckland, the area indicated by a red circle on the map alongside.

Curlew problems

The landscape changes associated with declining numbers of British breeding Curlew were reviewed in a 2017 paper by Samantha Franks and colleagues and summarised in the WaderTales blog Curlews can’t wait for a treatment plan. In 2020, Aonghais Cook et al analysed annual survival rates of Curlews in Britain and concluded that these were remarkably high, strongly suggesting that productivity is the main problem, as discussed in More Curlew chicks needed. Harry Ewing, a PhD student at the University of East Anglia, has been studying Curlews breeding in Breckland since 2019, in the hope that he might find answers to the question “Are there circumstances in which England’s lowland Curlew breed successfully?” The answer has implications for Curlews in other areas.

Targeting help for Curlew

Visitors to UK wetland nature reserves are becoming used to seeing fenced areas, in which species such as Avocet and Lapwing can nest relatively successfully, out of the reach of mammalian predators. These fences tend to shine a spotlight on the role of mammalian predation in wader declines but this is only part of the problem.

Landscapes used by ground-nesting birds have been massively changed by agricultural intensification and afforestation, which may make nests more vulnerable to predation. Additionally, human-induced climate change is rapidly altering environmental conditions. It’s hard to hide a nest if the grass won’t grow and difficult for chicks to find insects in a drought.

A set of tools is available to those managing land for semi-colonial waders in lowland wet grasslands (see Tool-kit for wader conservation). For Curlew breeding at much lower densities across human-modified landscapes, such as farmland, conservation is more challenging. In his IBIS paper, Harry Ewing quantifies variation in Curlew nest survival, in order to explore how management could be targeted to boost this key component of breeding productivity. Up to 80 pairs of Curlew were monitored annually between 2019 and 2021, in eight locations across Breckland, eastern England, an area in which nesting densities ranged from less than one to more than seven pairs per km2.

Breckland Curlew

The Brecklands of East Anglia have been shaped by humans for millennia. In their current form they comprise intensive farmland, commercial forestry plantations, and military bases and training areas, with pockets of land managed for the distinctive flora and fauna communities that used to be widespread across dry, grazed heath and grasslands. Curlew in Breckland can be found nesting alongside runways, within sugar-beet fields and on nature-reserves.

Curlew nest on an RAF base

In this study, pairs were monitored across a range of sites and habitats, including arable farmland, grassland, cleared forest, a military training area and an RAF base. The eight study sites were dominated by arable fields or semi-natural grassland, which is maintained by mowing or livestock grazing. Public access was restricted and some areas within grassland sites were enclosed by fencing. None of the fences were designed to protect breeding waders but their presence around military bases and to reduce grazing pressure in forests potentially provide benefits for Curlew, by reducing fox encounters.

Most of the fieldwork was conducted in 2019 and 2021, when 55 and 69 nesting attempts were followed. Curlew nests can be hard work to locate, and anyone who has ever studied breeding Curlew will be impressed by these numbers! The 2020 season started late, due to Covid-19 restrictions, but added another 12 nests.

The location of each nest was recorded with a GPS device, an iButton temperature logger was deployed in the nest lining, in order to capture the timing of nest failures, and hatching dates were predicted from the weights and measurements of eggs (more details in the paper). Regular monitoring and more frequent visits close to hatch date provided information on nest outcomes. Nest concealment was recorded on the day on which each nest was located, by measuring the height of vegetation around the nest.

Sugar-beet provides good cover, as the plants start to grow

Assessing nesting success

Harry Ewing, measuring an egg

Curlew breed in a wide range of densities across the Breckland study area. Six sites held between 0.17 and 0.72 pairs per km2, while two other hosted densities of between 3.3 and 7.4 pairs per km2. Nearly half of the probable or confirmed breeding pairs monitored annually were on the latter two sites, despite the fact that between them they only accounted for 5% of the study area.

A total of 136 Curlew nests were monitored across Breckland over the three summers, with the majority being found in unfenced grassland areas. 185 chicks hatched from 52 nests and 84 nests failed. Predation accounted for 86% of nest failures, with three nests failing during laying, one being trampled by cattle and five being lost during farming activities.

The key result in the paper is that hatching success was not high enough, in any habitat, in any year. There was some variation between sites but the mean probability of surviving incubation, for all hatched or predated nests was about 0.25, which is well below the estimated requirement for sustainability of a population.

Nest survival was similar in unfenced grassland, fenced grassland, arable fields and ground-disturbance plots within grassland, and across levels of nest concealment, so the spatial variation in nest survival was not the result of variation in management conditions or of nest concealment between sites. Only nine nests were found in fenced areas, making it unlikely that any positive effect of fencing could have been detected.

For nests for which data from temperature-loggers were available, it was possible to infer that 36 out of 44 predation events took place during the (short) nocturnal period, the time when foxes are most active.

Implications for Curlew conservation

Nest in disturbed grassland

It would have been great if Harry had been able to find a prescription for Curlew nesting success, by identifying a set of circumstances that delivered plenty of chicks. The fact that this was not possible, despite monitoring a large number of pairs, in a variety of habitats and across a broad density gradient, emphasises how hard it is going to be to preserve Curlew as a breeding species in England. Head-starting (raising chicks in captivity) may be able to give a temporary boost to numbers (see Will head-starting work for Curlew?) but these new recruits need to be able to breed successfully if local populations are going to be sustainable.

As the great majority (86%) of nests failed because of predation, boosting hatching rates of Curlew nests is likely to require actions to reduce mammalian predator impact across large areas of Breckland. It is much harder to help thinly-spread Curlew than semi-colonial Lapwing.

A conservation tool that is commonly deployed to reduce predation on ground-nesting birds’ nests is predator-exclusion fencing. Fences have the potential to double wader hatching rates in species such as Lapwing (Malpas et al. 2013) and it seems likely that they could also be effective at boosting Curlew nest survival, if provided at an appropriate scale.

Nest on a disused runway

In Breckland, fences were not deployed to protect Curlew nests  A small number of pairs nested within fenced areas but too few to detect an observable effect of fencing on nest survival. Increasing the number of nests enclosed by predator fencing in Breckland could potentially be achieved by deploying temporary electric fencing to protect individual nests but the substantial annual efforts required to locate nests and erect and maintain fences make this impractical, especially in areas where Curlew breed at very low densities.

The authors suggest that targeted deployment of fencing could potentially be effective if focused on ground-disturbance plots within grassland (see graphic below). These areas were originally developed for Stone-curlew but are often used by nesting Curlew, as described in a paper by Natalie Zielonka et al and summarised in Curlews and foxes in East Anglia. Plot-level fencing could be deployed at the start of the season, without the need to locate nests, and may benefit other ground-nesting species such as Lapwing. Unfortunately, these plots currently only support between four and seven breeding pairs of Curlew in the study area, suggesting only a modest potential increase in chick production.

Another alternative might be to erect permanent barrier fences along the boundaries of sites supporting high densities of nesting Curlew. Fencing the combined boundary length of the eight Breckland study sites that between them support about 80 pairs would require 185 km of fencing, which would be prohibitively expensive. However, given that nearly half of these pairs were breeding in just two of those sites, with a combined boundary length of only 14 km, perhaps this is where efforts might be focused? Assuming such fences would be as effective as for other waders, enclosing these two high-density sites with permanent barrier fencing could potentially boost the total number of chicks hatched in Breckland by about 90 chicks per year. It would be interesting to calculate whether this might be a more cost-effective way of producing fledged chicks than head-starting.

To sustainably maintain and recover Curlew populations in the wider landscape, in Breckland and elsewhere across the breeding range, actions outwith fenced areas are likely to be required. Lethal control of foxes, the main mammalian nest predator in the region, occurs across much of the Breckland study area and it would be useful to know how effective it is at limiting nest predation levels. This means learning more about predation, predator behaviour and control measures.

Across England, regional stakeholder networks are going to be important. Curlew territories are large and pairs move between habitats, implying a need to integrate evidence-based, Curlew-friendly policies into large-scale agri-environment schemes. Harry Ewing and his co-authors suggest that the next steps are to work with stakeholders to trial management actions for Curlew (e.g., fencing, head-starting and predator control). Actions to boost nest survival will need to be targeted in areas that contain suitable habitat to support chick growth and survival. Further research is required to understand the land management actions that can create and maintain such conditions at different scales.

To find out more

This blog focuses on the first paper from Harry Ewing’s thesis, undertaken by the University of East Anglia on a Natural Environment Research Council CASE studentship, as part of the EnvEast DTP. It was part funded by the British Trust of Ornithology, with support from RSPB and via a Defence Infrastructure Organisation Environmental Stewardship grant.

Nest survival of threatened Eurasian Curlew (Numenius arquata) breeding at low densities across a human-modified landscape. Harry Ewing, Samantha Franks, Jennifer Smart, Niall Burton & Jennifer A Gill. IBIS. doi.org/10.1111/ibi.13180


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

A Whimbrel’s year

There’s a lot to fit into twelve months if you’re a Whimbrel. In the last paper from his PhD, Camilo Carneiro assesses whether Icelandic Whimbrel can always manage to complete the annual cycle of migrate-breed-fatten-migrate-moult-fatten in just 365 days.

YY-LL (Yellow yellow – lime lime) collected data for several years.

What happens if a pair of Whimbrel loses a first clutch and a successful, late breeding attempt delays departure from Iceland, for instance? Is there the flexibility to make up for lost time during a west-African winter? Some answers are provided in a paper by Camilo Carneiro et al in The American Naturalist.

Facing the consequences

We know that large waders sometimes take a year off from breeding, as discussed in Teenage waders, suggesting that they may not always have the resources they need to breed every year. This blog, based on a paper by the Bird Ecology Lab team in Chile, tells the tale of a few Hudsonian Godwits that headed for the pampa grasslands of Argentina in spring, instead of migrating to Alaska.

As discussed in Gap years for sandpipers, taking a year off may make no difference to expected lifetime breeding output and could be more common when individuals spend the non-breeding season a long way from nesting areas or in poor quality sites. If Iceland’s Whimbrel are able to compensate for any delays they face during the course of the annual cycle then perhaps that suggests that all is well for this population that migrates all the way to West Africa at the end of the summer breeding season?

The story so far

Whimbrel only spend three months of the year in Iceland, with the rest of the time spent some 6000 km further south. In July or August, an adult will exchange the sparsely vegetated river plains and night-time frosts of Iceland for mud, mangroves and tropical temperatures. The diet changes too, from terrestrial invertebrates and crowberry to crabs.

Camilo Carneiro started a PhD on Iceland’s breeding Whimbrel in 2015, continuing the work of his supervisors, Tómas Gunnarsson of the University of Iceland and José Alves from the University of Aveiro (Portugal). Using geolocator tags he investigated the capacity for Whimbrels to undertake non-stop journeys, demonstrating that autumn migration was generally direct but spring migration for most birds included a stop, often in Ireland or the UK. You can read more in Iceland to Africa, non-stop. Birds that need to (or choose to) take a break delay their arrival in Iceland by about ten days.

Subsequent papers by the same team have shown that the most consistent point of the annual migration story is departure from Africa and shown links between weather and phenology. These two papers have been covered in the WaderTales blogs Whimbrel: time to leave and A Rhapsody of Whimbrel. In an attempt to discover the best places to spend the winter months, Camilo analysed tag and colour-ring data to work out links between conditions experienced in wintering locations and subsequent breeding success, as discussed in the blog Winter conditions for Whimbrel.

Seven years of data

To investigate how migratory animals navigate their annual schedule, and where and when they can make adjustments to their timings, Camilo Carneiro and his colleagues used annual-cycle data of 38 Icelandic whimbrels tracked over 7 years. They asked three questions:

  • Does the change in the timing of one event in the annual calendar, such as a late breeding season, affect the timing of subsequent events, perhaps with further down-stream domino effects?
  • Can individuals compensate for delays, on migration for instance, by spending less time on the next stage of the annual cycle, e.g. by reducing a stop-over?
  • Are there potential fitness consequences? Do birds that are subject to delays breed later? In waders, earlier chicks are more likely to recruit to the breeding population so being just a few days late returning to Iceland may have consequences.

During the period 2012 to 2018, a total of 78 geolocators were deployed on Whimbrel breeding in Southern Iceland. The device was attached to a leg-flag in one year (see picture) and usually collected in the subsequent breeding season. In most cases, a replacement geolocator was fitted, in order to collect further data on that individual. Unsurprisingly, it became harder to catch tagged birds over time, as birds learned to recognise nest traps and research vehicles. The fact that so much valuable information was collected from the same birds is testament to Camilo’s patience. Sixty-six geolocators were retrieved from 39 individuals. Birds could only be caught when incubating a full clutch of eggs so nest losses due to predation affected the likelihood of recapture.

Please see the paper in The American Naturalist for full details of the methods used to collect data on breeding success and for interpretation of data collected using geolocators.

Whimbrel YY-LL

Before looking at the results in the paper, here’s an example of data collected using geolocators, for two years in the life of YY-LL (Yellow Yellow – Lime Lime), pictured alongside.

In 2015, YY-LL nested successfully and left Iceland on 16th August. After four days of direct flight, he reached Guinea-Conakry. His nesting attempt in 2017 was unsuccessful and he migrated south a little earlier, on 5th August, again taking four days to fly 6000 km.

In the springs of both 2016 and 2018, YY-LL left the winter grounds on 22nd April, arriving in Ireland on 26th April in 2016 and on 25th April in 2018. Migration from Ireland to Iceland took place between 5th and 7th May in 2016 and between 7th May and 9th May in 2018.

YY-LL hints at strong consistency of spring migration timing, independent of nest success.

Results

The 38 tagged Whimbrel provided information about 76 autumn migrations and 60 spring migrations. Most of the birds (89%) spent the winter between Senegal and Sierra Leone, particularly in Guinea-Bissau and Guinea, with one bird in Portugal and the rest in coastal northwest Africa.

As is usual in waders, females left Iceland before males, the difference being typically around six days. Although failed breeders did depart earlier than birds that successfully reared chicks, the difference was not great, again averaging around six days.

There was no suggestion that birds that left Iceland late in the season ended up on a delayed schedule for return to Iceland in the subsequent spring. This implies that resources in Africa were sufficient to ‘catch up’ with earlier birds, despite the need to complete a full moult and to prepare for another 6000 km migration.

As indicated in previous papers (and in the blogs mentioned above) spring departure dates of tagged birds from Africa were not different for different countries or for different sexes. However, birds that stopped in Europe tended to leave Africa earlier (19th April on average) than those that made direct spring flights (30th April). These latter birds tended to arrive in Iceland about a week earlier than birds on a two-stage migration, representing a neat, overtake manoeuvre! Typically, males arrived in Iceland about a week earlier than females.

For the sample of tagged birds, neither the autumn departure date from Iceland nor wintering location had any apparent effect on the arrival date in the next spring.

There seems to be a strong signal that, just as YY-LL did, an individual Whimbrel can make up for any delays incurred, with birds that arrive at a location later spending less time there, whether that be a wintering site or a spring stopover location. Females tended to spend longer at spring stopover locations, which ties in with the earlier arrival of males in Iceland. The graph alongside illustrates these two points. Birds that left Africa earliest spent more than 15 days at stopover sites but birds on a later schedule stopped off for as few as 6 days. Triangles represent males and squares are females.

Over the course of a year

Putting this all together, Camilo and his colleagues found that individuals appear to use the wintering sites to compensate for delays, these mostly having been associated with a successful previous breeding season. The wintering season is up to 38 weeks long so a Whimbrel that heads south a little late has plenty of time in which to catch up with earlier birds. The timings for other wader populations that spend shorter periods in wintering locations may be more constrained, given that post-breeding moult might take 20 weeks and the time to fatten up for migration can add an extra seven weeks.

Once a bird leaves Africa, it is harder to compensate for delays, although attempts are made to do so, with later individuals stopping for shorter times at spring stop-over sites and then nesting shortly after arrival in Iceland. Data in the paper suggest that it is not possible to catch up completely, before the start of the breeding season, if time is lost on the way north. Given the known link between lay-date and nesting success, these spring delays may have consequences for productivity and reduce the capacity to re-nest following clutch loss.

Iceland’s Whimbrel

Camilo’s research suggests that adult Whimbrel in the south of Iceland have the capacity to make up for any delays that they face during the annual cycle. The same may not be true for other large waders, the populations of which are mostly in decline. The blog Why are we losing out large waders? reminds us that two curlew species are thought to be extinct or on the verge of extinction and that most of the rest are in trouble.

There are increasing pressures on Whimbrel breeding in Southern Iceland, associated with new forestry plantations and infrastructures such as road developments and power lines

All is not necessarily well for Iceland’s waders either. Two 2022 papers by Aldís Erna Pálsdottir, looking at the effects of forestry and power-lines, suggest that the Whimbrel is one of the wader species most seriously impacted by ongoing changes to the Icelandic landscape, as discussed in Power-lines and breeding waders and Iceland’s waders need a strategic forestry plan. These pressures seem to be reflected in poorer breeding output, as suggested by counts of family parties of Whimbrel made by Tómas Gunnarsson and colleagues in annual June and July surveys.

Camilo’s paper may indicate that adult Whimbrel can cope with all that life throws at them but if they cannot raise enough chicks the species will still be in trouble. With financial support from the Icelandic Centre for Research, Camilo is now studying the challenges that chicks face, as they prepare for their first migration from Iceland to Africa.

Here’s the link to the paper:

Annual schedule adjustment by a long-distance migratory bird. Camilo Carneiro, Tómas G. Gunnarsson & José A. Alves. The American Naturalist. doi.org/10.1086/722566


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

Will head-starting work for Curlew?

83 captive-reared Curlew were released successfully in 2019, over 130 in 2021 and a similar number in 2022 but this does not mean that head-starting is a solution to England’s Curlew problems. We don’t yet know the proportion of youngsters that survive the difficult ‘teenage years’, how many will find suitable breeding habitat and whether these birds can reliably raise chicks of their own. What have birds such as 7K, pictured here, revealed and what might happen next?

Exciting news

There has been some great TV and press coverage of head-started Curlew this summer (2022), with films of captive-reared birds being released from pens and flying for the first time. But what does the future hold for naïve birds that take to the air in Shropshire & Powys, the Severn & Avon Valleys of western England, Cornwall, Sussex and Norfolk? The first signs are good, as illustrated below in maps published by the BTO for the birds that they are tracking in Norfolk.

Satellite-tagged birds released at Ken Hill (Black stars – close to the Norfolk coast) can be seen flipping backwards and forwards between the mudflats and their release site, before beginning to explore more of the Wash estuary.

Birds released further inland, on The Sandringham Estate (White stars), started foraging in grassland areas and it took a while until any of these birds discover the coast. One bird “7Y” moved onto a little-used airfield (below), where it was seen with a flock of adults. It appears to have adopted a mudflat/grassland tidal routine, involving ‘commutes’ of ten miles each way, possibly having been guided by the adults (top row, third from the left).

One of the Sandingrham youngsters joined up with a flock of adults on an airfield five miles from the release site

It is easy to get ‘wowed’ by the TV reports and the individual stories of tracked birds but these are early days for Curlew head-starting. There’s a lot still to learn!

Why head-start Curlew chicks?

This Curlew is nearly ready to fledge

The Eurasian Curlew is in trouble, as you can read in the WaderTales blog “Is the Curlew really near-threatened?”. Annual survival rates of adults are high but productivity is low, with an estimate of 10,000 too few youngsters being fledged each year, to maintain population levels in the UK. The Irish Curlew population has crashed, providing dire warnings of what may happen soon in Wales, where there has been a decline of 73% since 1995 (Breeding Bird Survey). In southern England, most of the losses happened before 1995 and only a few populations remain, hanging on in areas such as Breckland in East Anglia.

Lessons from Black-tailed Godwits

We know that head-starting works well for limosa Black-tailed Godwits breeding in East Anglia but there was no guarantee that this would be the case when the WaderTales blog Special Black-tailed Godwits was published in 2017, to coincide with the departure of captive-reared birds from their WWT Welney release site. When the first birds not only returned in 2018 but actually raised their first chicks (see Head-starting Success), the approach looked to be successful. Over the following years, with more releases, the godwit population in the Ouse and Nene Washes has increased significantly. This temporary relief package for a beleaguered population has worked well, providing time to try to improve breeding conditions in the wild, so that Black-tailed Godwits can look after themselves. Head-starting is an intervention of last resort and cannot be a long-term solution to the problems faced by the species.

Young Curlew is now identified as 2X

It is possible to support local populations of scarce birds, such as Bitterns and Little Terns, with interventions such as increasing areas of suitable habitat and protecting nest sites, but doing the same for Curlew will be more of a problem. Here, the challenge is to halt a decline of a species that breeds across wide areas of the UK, mostly on farmed land that is not being managed for conservation. Intensive, local action is not going to be enough. This is going to need teamwork between landowners, conservation organisations and volunteers – acting at a scale that has not been tried previously. Perhaps head-starting will help?

A pragmatic plan

The most recent UK population assessment of Curlew, in British Birds, suggested that there were 58,500 breeding pairs in 2016. Numbers have almost certainly dropped since then but the species is nowhere near as threatened as England’s breeding Black-tailed Godwits. The biggest Curlew head-starting programme, led and funded by Natural England, was a response to an opportunity to use unwanted Curlew eggs – not a ‘last-chance’ solution, as it had been for Black-tailed Godwits. Each year, Curlew eggs were being collected on RAF airfields, under licence, to deter adults and reduce airstrike risk. Thanks in no small part to Natural England’s Graham Irving, many of these eggs are now being head-started, instead of destroyed.

What happens to head-started birds?

Curlew eggs in an incubator

The Natural England project builds upon the experiences of the Curlew Country team, working in the Shropshire hills and Powys borders. They released 6 head-started Curlew in 2017, 21 in 2018, 33 in 2019 and 34 in 2021. Raising extra chicks is part of an initiative that involves a broad range of stakeholders. See their website for more information.

The first stage of the head-starting process works well. Aviculturalists can rear and release chicks, with very high rates of success. They are learning more and more about the best diets, appropriate husbandry and the release process. Individual chicks wear small colour-rings, so that progress can be monitored daily, and they receive leg-flags and get weighed and measured before release. For instance, the young Curlew 7K featured above is a GPS tagged male that weighed in at a respectable 550 grammes when it was transferred to its release site on The Sandringham Estate, on 14 July 2022. By the end of August, it had moved to the Wash Estuary, as you can see in the figure above (top left map). There was a December report of 7K from a piece of amenity land on the edge of a new housing development in Hunstanton.

Before transfer to The Sandringham Estate release site, a young Curlew is checked over by a vet

From the start, the Natural England project has been built on partnerships. In the first year (2019), eggs were hatched and chicks were reared at WWT Slimbridge. Fifty young Curlew were released near-by and the first of these birds (wearing ring number 23) was found nesting near Gloucester in 2021, raising one chick of his own. In 2022, three years after release, five of the head-started Curlew were on breeding sites in the Severn and Avon area and one bird had moved to the Thames Vale. Male “23” shows that head-starting can add more Curlews to the breeding population but is he exceptional? What proportion of released birds make it this far and go on to raise chicks? The video by Kane Brides in this WWT blog tells the West Country story so far.

After a Covid-caused hiatus in 2020, the head-starting project expanded in 2021, with birds reared at both WWT Slimbridge in Gloucestershire and Pensthorpe in Norfolk. Richard Saunders, the Senior Ornithologist for Natural England, recognised the importance of learning what happens to newly-released fledglings and of monitoring potential recruitment. Working with Sam Franks of BTO and with lots of support from others, the Pensthorpe-reared Curlews are starting to reveal some of their secrets.

Where will the Pensthorpe birds set up home?

Most waders are philopatric; when they look for breeding sites, they tend to return to places close to the places in which they were raised. The release sites in North Norfolk were chosen not only because they provided suitable conditions for growing teenage waders, with low predator numbers and adults feeding nearby, but also because they are relatively close to current breeding sites, such as Roydon Common and the heaths of Breckland (see Curlews and foxes in East Anglia). There are also airfields, of course, and it will be unfortunate if birds choose these sites, given that eggs were removed to try to reduce bird-strike risk.

Radio tag (left) will drop off when juvenile feathers are replaced. Satellite tag is mounted on a temporary harness and worn like a rucsac.

All of the chicks are colour-ringed, some of them are radio-tagged and others are GPS-tagged, prior to release. By following tagged birds, as they explore the area around the release sites, the project aims to understand more about habitat use and to see if these naïve birds seem unduly prone to predation, given that they have not been trained to look out for danger by watchful parents. GPS tagging helps to paint the bigger picture; would birds move to the mudflats of the Wash and spend the whole winter in Norfolk or would some move on, to southern England, Ireland or France, for instance? These data will hopefully be augmented by reports of colour-ringed birds that do not carry tags.

2C takes to the air after release at Ken Hill

The first results have been encouraging. For example, a GPS tagged female from the first cohort released in Norfolk in 2021, wearing flag 0E, has followed the ‘stay local’ option. She spent the winter on the saltmarshes of RSPB Frampton, on the Wash in Lincolnshire, and has largely stayed on the south shore of the Wash through her first summer, showing no evidence of visiting any type of breeding habitat yet. A male Curlew wearing flag 4P has been more adventurous, spending the winter on the Exe Estuary in Devon, a site used by some adult Curlews that breed in Eastern England.

The amazing left-hand map below shows the route taken by ‘6Y’, one of the first batch of 2022 birds to be released on The Sandringham Estate. The story of this bird is told by the BTO’s Dr Sam Franks:

“The first of this year’s birds to migrate away from Norfolk departed at sunset last Wednesday & arrived on a Staffordshire field at sunrise on Thursday. It then flew towards Ireland & made an anxiety-inducing trip out into the Atlantic before returning to dry land.”

The right-hand map shows an overnight flight by ‘9L’, one of the last birds to be released from Ken Hill. This bird, pictured to the right , set off on the evening of 16th September, flew southwest overnight and headed south when it ran out of land. It landed in France at 01.30 on 17th. Although there is supposed to be a Curlew hunting ban in France (see this blog), three satellite-tagged birds from elsewhere in Europe were shot in the first weekend of the 2022 autumn hunting season. Will 9L survive the winter?

Data generated by colour-ring sightings will be analysed to check whether annual survival of youngsters in their first couple of years are consistent with figures for wild-reared chicks. This follow-up work is really important – the head-starting operation may seem to be successful but if few chicks survive long enough to breed then alternative approaches may be needed. Sightings of colour-ringed birds provide important information to add to data collected from tagged birds, which means that there is a vital role to be played by birdwatchers.

Around England

People care deeply about Curlew, as Mary Colwell explored in a recent book, reviewed in the WaderTales blog Curlew Moon, so it is not surprising that landowners feel inspired to help the species, by releasing birds on their own land. An attempt is being made to boost a tiny population on Duchy of Cornwall land on Dartmoor, using eggs from East Anglian airfields that were hatched at Slimbridge, and a licence has been granted to take eggs from a site in Northern England to be reared in Sussex, on an estate that appears suitable but where Curlew do not currently breed.

Release sites for these translocated birds do not hold many (or even any) Curlew so it will be interesting to learn whether the behaviour patterns of these fledglings are different to those of birds released in Norfolk and in the Severn and Avon Valleys of western England. All birds are being ringed, by WWT, GWCT and BTO ringers, with every report of a marked bird adding to our understanding of the success of the various projects. In each case, hopefully there will be enough money to deploy dedicated fieldworkers to monitor what happens to the released birds during the crucial first few weeks of independence, so that fledging rates can be accurately assessed, and to monitor return rates in subsequent breeding seasons. For slow-maturing birds, this follow-up work will involve a five-year commitment.

Maximising effectiveness

The work being funded by Natural England in East Anglia is expensive. It will be judged as successful if released birds augment local populations, whether these be in East Anglia (as hoped) or in another part of lowland England. If birds choose to breed in the North of England, where numbers are still high, or even overseas, then that may make it difficult to justify the expense of further rearing and monitoring work.

There is a concern that head-starting will be seen as a solution to the problems being faced by Curlew. It isn’t! The estimated shortfall in fledged chicks is 10,000 birds per year, across the whole of the UK, and head-starting will never make a big impact on that number. It may be a way to boost numbers in lowland areas, from which the species would otherwise be lost, but only if the conditions for successful breeding can be created and maintained. This means tackling the thorny problems of habitat degradation and predator numbers. Head-starting may seem like a dynamic intervention but if birds are released into areas where breeding success is too low then it’s not going to produce a sustainable solution to the problems being faced by England’s Curlew.

Photos of ringed birds are particularly appreciated

The Norfolk head-starting project would appear to tick all the right boxes – release sites with low predator numbers, right next to an estuary with lots of Curlew and with successful breeding sites near-by. Head-started birds soon start to explore the mixture of arable and tidal resources available to them and it looks as if early-years survival rates might be as high as expected of wild-reared birds. Despite all these positive signs, it will be a couple of years until this year’s young Curlew start breeding – somewhere – and that will be the crucial test of head-starting. Meanwhile, it is hoped that birdwatchers will look out for colour-ringed birds, so that survival rates can continue to be monitored. Every Curlew counts!

Norfolk-ringed head-started Curlew wear yellow flags, each with number & letter, immediately above an orange ring. Please report sightings of colour-ringed head-started Curlew using THIS LINK.


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.

When mates behave differently

Iceland’s 40,000 Oystercatchers are an interesting mix of resident birds and migrants, providing an ideal system in which to study the costs and benefits of the two options, and to try to work out what influences whether an individual becomes a ‘resident’ or a ‘migrant’. I’ve added the inverted commas because many residents migrate within Iceland in spring and autumn; it’s just that they don’t undertake long-distance flights across the Atlantic.

In their paper in Ecology & Evolution, Verónica Méndez and colleagues from the South Iceland Research Centre (University of Iceland), the University of Aveiro (Portugal) and the University of East Anglia (UK) investigate the timing and success of breeding attempts by resident, migratory and mixed (resident/migratory) pairs of Icelandic Oystercatchers.

Iceland’s Oystercatchers

As outlined in Mission Impossible: counting Iceland’s wintering Oystercatchers, about 30% of Icelandic Oystercatchers never leave the country, coping with cold temperatures, short December and January days and a restricted diet. In the winter months they can be found in the tidal zone of a few estuaries, mostly in the warmer west.

The majority of Iceland’s Oystercatchers fly 1000 km or more across the Atlantic, to Ireland, the UK and the coastal fringe of western Europe. Here, many colour-ringed birds have been spotted by birdwatchers, who play a vital part in migration studies. The blog Which Icelandic Oystercatchers cross the Atlantic? revealed that males and females were equally likely to migrate, while there appeared to be no assortative mating in spring (residents don’t preferentially choose resident partners, for instance).

It would be easy to envisage circumstances in which resident birds might be at an advantage, at the start of the spring breeding season, having not had to cross the Atlantic and thus being ready if an early nesting opportunity opens up. On the other hand, in a cold spring or after a particularly harsh winter, perhaps they could be in poorer condition than newly arrived migrants, and this may potentially delay breeding. What actually happens?

Fieldwork

Both resident and migrant Oystercatchers breed throughout lowland Iceland. Within breeding pairs, it is estimated that about 20% of pairs are resident, 46% are migrant and 34% are mixed. These are long-lived birds that generally maintain the same partners between years, despite the fact that individual males and females may spend seven months of the year up to 3000 km apart. Parents tend to be equally involved in incubation duties, territorial defence and chick rearing, although males tend to remain with their youngsters longer than do females.

Between 2015 and 2018, Verónica Méndez and her colleagues monitored the breeding attempts of Oystercatchers in southern Iceland, continuing a study of marked individuals that started in 2013. Adults were caught on the nest and sexed by later analysis of feather samples. With the help of a network of volunteer observers, the winter locations of 186 (out of 537) marked birds had been established when the paper was first written. Using these known outcomes and with additional information from stable isotope analysis, it was possible to assign the remaining 351 birds as ‘residents’ or ‘migrants’. Amazingly, 73 of these 351 birds have been seen since the isotope data were analysed and all of the assumptions on winter locations were found to have been correct.

Early nesting attempts may be hampered by spring snowfall

The first migrant Oystercatchers arrive in Iceland in February but no nesting has been recorded before mid-April. Searches for colour-ringed birds and nests were conducted every 2-3 days and then nests were followed through to hatching or failure. Second (and third) nesting attempts were also monitored. Oystercatchers remain in the vicinity of the nest after hatching their chicks and then feed them throughout the growing period. Chicks were metal-ringed just after hatching and individually marked with colour-rings when around two weeks old. Families were monitored every 3-4 days until all chicks were fledged or lost, allowing productivity (number of chicks fledged per pair) and fledging success (number of chicks fledged in nests where at least one egg hatched) to be recorded.

Who breeds when?

Verónica and her colleagues were able to estimate laying dates for 138 pairs with known migratory behaviour (56 migrant, 50 mixed and 32 resident pairs) in one or more seasons during 2015-2018, providing a total of 228 observations.

The top graph shows that, on average, 2015 was a much later breeding year than the other three. This was a colder spring; the sort of colder conditions that an older Oystercatcher may well have encountered frequently in its youth! (The longevity record for BTO-ringed Oystercatcher is 41 years – see Waders are long-lived birds – and the trend for there to be more frequent warmer springs is discussed in this Black-tailed Godwit blog).

The lower graph shows a breakdown of the data into the three categories – Resident (black dots), Mixed (grey) and Migrant (white). There is no difference between the egg-laying dates for residents across the four years. However, in the 2015 breeding season, in cases where either member of the pair is a migrant, there was an average nesting delay of over a week. An analysis in the paper shows that it does not matter which member of a mixed pair was the migrant, the delay in 2015 was the same.

Reproductive performance

Unusually amongst waders, adult Oystercatchers feed their chicks

As expected, Oystercatcher pairs that made earlier nesting attempts were more likely to lay a replacement clutch after nest loss, had higher productivity and higher fledging success. This is in line with the modelling paper described in Time to nest again. Early-nesters tended to have bigger clutches too. Any differences between the performance of residents, mixed pairs and migrants could be accounted for just by the timing of nest initiation.

In the papers’ Discussion, the authors suggest that, in the three warmer years, earlier nesting of pairs that included at least one migrant was sufficient to slightly enhance nest success but not overall productivity, above that achieved by pairs with residents. The migratory behaviour of the male within a pair appeared to have a stronger effect on fledging success than the migratory behaviour of the female, suggesting that males may play a more important role than females at the chick stage. This is interesting in the context of previously-published research by Verónica and her colleagues, as described in The Dad Effect blog.

What does this all mean?

In other studies, described in the Discussion, residents in systems where some individuals migrate have been found to have advantages over migrants, because they can get on with breeding earlier. This was not the case for Icelandic Oystercatchers, potentially because migrants can arrive in good condition in all but the coldest of years.

Hatching brood of three

In the cold year of 2015, Oystercatcher pairs nested an average of between a week and 12 days later than in other years. This delayed nesting occurred in migrant and mixed pairs but not in resident pairs, suggesting that the effect of the severe weather may have been greater on migrants than residents. Cold spring conditions in Iceland tend to be part of a wider pattern of cold weather across northwest Europe. The authors suggest that wintering conditions might influence the body condition required to reproduce and that these conditions may be more variable for migrants.

Only one cold year occurred during this study, so the authors don’t know whether pairs with migrants consistently breed later in colder years. Given that cold springs are increasingly rare in Iceland, 2015 may turn out to have been one of the few remaining opportunities to reveal the dynamic nature of links between weather, migratory behaviour and breeding phenology at these latitudes.

One potential explanation of the difference in the timing of nesting is the effect of habitat. The Icelandic team has found that there is a strong tendency for migrants to breed inland, whereas residents tend to breed along the coast. During the cold spring of 2015, inland habitats were not available as early as in the following years (everything was frozen), mostly delaying the breeding attempts of migrant and mixed pairs, rather than residents pairs.

Long-term studies

Verónica Méndez with one of the marked birds

The take-home message of the paper by Verónica Méndez and her colleagues is that it pays to nest early, which is not unexpected. Perhaps it is surprising that, in the cold spring of 2015, mixed pairs still bred at the same time as pairs of migrants, suggesting that residents waited for their migrant partners. Perhaps, the benefits of nesting with the same partner are very strong, or finding an alternative mate is difficult or both?

The study suggests that the links between individual migratory behaviour and reproductive success can vary over time and, to a much lesser extent, with mate migratory behaviour. Understanding these effects of pair phenology on breeding success may help researchers to understand the potential impacts of changing environmental conditions on migratory species. Such variation is very difficult to capture unless long-term funding is available. Four years may seem like a long time to observe the same Oystercatchers but, for birds that may easily live twenty years, this is nothing!

The full paper can be found here:

Effects of pair migratory behaviour on breeding phenology and success in a partially migratory shorebird population. Méndez V., Alves J.A., Gill, J.A., Þórisson, B., Carneiro, C., Pálsdóttir, A.E., Vignisson, S.R. and Gunnarsson, T.G. Ecology & Evolution


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

A Norfolk Curlew’s summer

This blog follows the fortunes of one Curlew (B/OW-OY) as he attempts to raise chicks in Norfolk, migrates to Portugal, diverts to avoid a forest fire and returns to his winter home in a Portuguese estuary. Using tagging data we know that he left his Norfolk breeding territory in England on 17 June 2022 and arrived at Montijo, in the Tagus Estuary of Portugal, on 16 July – but that’s only part of his story.

Curlew migration

B/OW-OY (named ‘Bowie’ in a BTO press release) may have been travelling between Norfolk and Portugal for a decade or more, flying backwards and forwards between the dry grassland of inland East Anglia and the coastal mud of the Tagus Estuary. We have new insights into his life because he was ringed and tagged on 2 April 2022, as part of a project to understand Curlew breeding success rates in East Anglia – about which more later!

Curlew migration is complex. The birds that we see in autumn and winter in the UK include some local birds but the majority will have crossed the North Sea, particularly from Finland. The map alongside is taken from the BTO website. As you can see, some British-bred Curlew migrate south for the winter. At the Europe-wide scale, the major axis of migration is also from northeast to southwest, as you can discover using the European African Bird Migration Atlas mapping tool. ‘Bowie’ may be feeding in Portugal alongside birds that have bred in Poland, Finland and France.

Breckland Curlews migrate in all directions in the autumn. At least three have been found wintering on the Norfolk coast, five colour-ringed birds have travelled to southwest England (Dorset, Devon and the Severn Estuary), another of the tagged birds has flown to Ireland and a colour-ringed bird has been seen in northern Spain.

Breckland Curlew

Harry Ewing is studying Curlew in Breckland – the tapestry of heath, grassland, forestry, arable farming and military bases in the dry, heart of East Anglia. His PhD at the University of East Anglia is being undertaken in partnership with BTO and with input from RSPB. Harry is working across a range of habitats that are used by nesting Curlew, from airfields to sugar beet fields. The breeding site of B/OW-OY is a grassland heath that is managed in ways that maintain habitats that are suitable for scarce plants, invertebrates and birds. Here, pairs of Curlew breed alongside Stone-curlews and Woodlarks.

Most of Harry’s Curlew struggle to fledge any chicks, as is the case in many places across Britain and Ireland, and it will be interesting to see the papers that come out of his PhD, in due course. Two previous WaderTales blogs describe the predation problems Curlews face in East Anglia (Curlews and foxes in East Anglia) and give information about conservation interventions that suit breeding waders (Grassland management for Stone-curlew).

Catching Curlew

During his PhD, Harry has found up to 70 Curlew nests in a single breeding season and followed their outcomes until the eggs or chicks were predated or through to fledging. Harry checks on his breeding Curlew on an almost daily basis but does not always see adults on territory. By catching birds at roost, at the start of spring, and deploying a small number of GPS tags, funded from the BTO Curlew Appeal and by Natural England, he hoped to learn more about how birds use surrounding habitats and to get more accurate information about the timing of breeding events. The extra data about migration that he gets from birds like B/OW-OY are a bonus.

Harry had been looking for opportunities to catch Curlew in the Brecks when they returned to the areas in the spring. A flock of roosting birds was spotted by local graziers on 31 March 2022 and, with the help of the Breckland Farmers Wildlife Network, Nigel Clark and the Wash Wader Research Group, a catch was made on the evening of 2 April. Eleven birds were caught, all of which were colour-ringed and eight of which were also fitted with GPS tags by Sam Franks of BTO (see picture).

A ‘miracle chick’

Harry spotted B/OW-OY on the day after capture, on a grass heath just 3.7 km away from the ringing site. As luck would have it, he was paired with B/OW-OL, a newly colour-ringed female. Harry could not download data from the tagged bird, due to a lack of signal coverage, but he saw OY and OL almost daily over the next two months. Information collected on OY’s tag and sent from Portugal in July filled in some of the gaps. OY fed mostly on cultivated plots (as described in Grassland management for Stone-curlew) but he also used nearby arable farmland, especially an asparagus field about 3 km from the heath, and made a brief trip to a local airfield during the pre-breeding period.

Both OY and OW were observed regularly and Harry found their nest and a completed clutch on 6 May. As can be seen in the photograph, it was in a very open site, on dry, grazed grass heathland. When Harry checked the nest on 25 May, three chicks had hatched and one was still hatching. A second check the next day, to confirm successful hatching of the final chick, revealed a sad tableau of overnight predation: one chick was dead, one was injured and two were missing. Injuries suggested a mustelid attack – probably a stoat (see photograph above). That was the end of the breeding attempt – or was it?

Another day later, on 27 May, Harry discovered a bird he named ‘miracle chick’; a one-day-old youngster that was feeding near the nest site, on its own, with no parents guarding it. OY and OL were still present on the heath but had joined a flock of failed breeders, presumably having assumed that their chicks had all perished. Two days later, ‘miracle chick’ had moved 100 metres and latched onto a family of two 25-day-old chicks, feeding with them and sharing the protection afforded by their parents. OY and OL were still in the post-breeding flock, presumably unaware that their breeding season had not been a total failure.

Curlews gather in small flocks when they lose their chicks or nests. For them the season is over.

As Harry continued his daily checks of broods around the Brecks, to try to understand more about the predation of chicks, he was able to establish that the first step-sibling of ‘miracle chick’ was predated on the night of 31 May/1 June and the second on the next night. Nocturnal predation of big chicks is probably by foxes. ‘Miracle chick’ continued to be defended by its step-parents until 16 June. On 17 June, Harry found an adult female with a broken wing and the chick was never seen again. Perhaps she was the step-mum and the injury was sustained in unsuccessful defence of ‘miracle chick’. ,We shall never know. By this time, OL and OY appeared to have left the heath; the last sighting of the female was on 10 June and the tagged male was not spotted after 16 June.

Heading south

Thanks to the information that was transmitted from Portugal, we know that B/OW-OY (Bowie) left Breckland at 21.30 (UTC) on 17 June, arriving on the Essex coast at Brightlingsea just 80 minutes (and about 50 miles) later. He spent the next four weeks feeding around the coast of Mersea Island, leaving at 20.00 on 14 July. There was a nine-hour lay-over in Spain on 15 July, before another evening departure (20.40). The last part of the journey is interesting; it included what looks like a diversion around the forest fires that were burning in the Ourém region in Portugal at that time (see map).

It is fascinating to think about how this one Curlew from Norfolk might have travelled to Portugal. Did it fly to the coast alone? Did it meet up with other birds that were heading south from Essex and fly in a flock? Did it ‘change flocks’ in Spain, as we would planes at an airport? This blog, based on a recent French paper, gives a feeling of what it is like to be a migrating Curlew: The flock now departing.

Threatened winter home

Bowie arrived on the Tagus Estuary at 09.30 on 16 July and he may well stay here until next February or even March. The Montijo peninsula, which appears to be his winter home, is one of the most important parts of the Tagus Estuary for Curlew. They feed in the mud and roost on salting islands, following the rhythm of the tides. This estuary is right next to Montijo Air Force Base.

The saltmarsh at Montijo on 18 July 2022. Josh Nightingale counted 22 Curlew.

Montijo Curlew are accustomed to the relatively infrequent take-offs and landings by military aircraft, in the same way that Curlew on RAF bases in East Anglia are happy to breed on the grassland alongside runways. All this will change, however, if the plan to expand the site, lengthen the runways and introduce all-day flights of passenger aircraft goes ahead (planned to operate at 5 minute intervals). Add on control measures to reduce the potential of bird-strikes to aircraft and Montijo and the Tagus as a whole will become much less attractive to hundreds of thousands of waterbirds. As you can read in Tagus Estuary: for birds or planes, we need to fight very hard to stop this new airport which is on a specially protected Ramsar site. If this jewel on the East Atlantic Flyway can be degraded then what is there to stop similar developments elsewhere?

One tagged bird

It looks as if B/OW-OY skirted around forest fires in the Ourém region.

In the spring, B/OW-OY will return to his Breckland breeding site, hopefully to meet up with B/OW-OL and to have another go at raising chicks. For him nothing has changed – he’s doing the same again for another year. The only difference is that he is tagged now, and that provides us with insights into his world and the challenges that he and other migratory waders face. Imagine what it will be like if, next year or in the near future, he arrives back in Montijo and finds bulldozers and freshly-poured concrete, instead of mud and saltmarsh. With a new airport to contend with and yet more anthropogenic change to his world, life does not look as if it will get easier.


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 12th 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 6th May and the 20th 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:


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

Navigating a vast ocean

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

Coping with wind

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

Migrating Hudsonian Godwits are heading for Alaska

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

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

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

Tracking transoceanic journeys

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

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

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

Results

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

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

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

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

Learning more

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

The full paper can be found here:

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

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

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


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

When Oystercatchers can’t find food

In a rapidly changing world, wintering waders face unprecedented challenges. How much flexibility is there for individuals to cope with issues such as over-fishing of shellfish stocks, habitat removal, pollution, and the effects of rapid climate warming on their food supplies?

Colour-ring records show that many wintering waders tend to be site-faithful, feeding in the same estuaries and even the same small patches year after year. This makes sense if food supplies are reliable and predictable but what happens when there is massive change in food abundance? In a 2021 paper in MEPS, Katharine Bowgen and co-authors describe the impacts of a cockle die-off in the Burry Inlet (part of the Severn Estuary in Wales) on the local population of Eurasian Oystercatchers. Their findings illustrate how important it is to protect networks of sites, rather than individual inlets or estuaries.

Assessing the options

When food supplies are low – or crash suddenly – what can birds such as Oystercatchers do? There are three likely options in these circumstances:

  1. Wait and hope
  2. Move elsewhere and never return
  3. Move elsewhere until conditions improve
When food availability is low, some Oystercatchers may not complete wing moult

If ‘wait and hope’ is the main option then, following an event that reduces food availability or abundance, there should be little evidence of birds dispersing, measures of annual survival might be depressed for a period and, once conditions improve in the area, young birds might be likely to fill the spaces available. This should lead to higher proportions of sub-adults in the period immediately after such a hiatus and a steady increase in numbers.

Movements to other sites, whether temporary or permanent ought to be detectable from mid-winter counts and through reports of ringed birds. The pattern after a permanent shift in birds would be similar to that following mortality; again, young birds might be expected to take advantage of food resources as they recover In reality, of course, individual birds that use a site may opt for any one of the three options, creating a mixed picture.

Oystercatcher numbers

Over recent decades, numbers of Eurasian Oystercatchers have declined. In 2015 the species was reclassified as “Near Threatened” on the IUCN’s Red List (Birdlife International) and “Vulnerable” within Europe. It is also Amber listed on the UK’s Birds of Conservation Concern list, due to its European status, the concentration of its wintering population in protected sites and the international importance of UK breeding and wintering populations.

Colour-ring sightings can improve survival estimates

The changes in Burry Inlet Oystercatcher numbers should be viewed within a pattern of national declines. Since the 1990s, winter numbers on UK estuaries have dropped by a third, back to levels seen in the 1970s. Patterns around Great Britain vary (see below); Welsh numbers on estuaries, as assessed by WeBS, have held up well but there has been a 25-year period of decline in England and there was a sudden fall in Scottish numbers at the start of this century.

The Burry Inlet

The intertidal mudflats of the Burry Inlet in south Wales are of international importance for non-breeding waterbirds of various species, including the Eurasian Oystercatcher. This SPA (Special Protection Area) is part of The Carmarthen Bay and Estuaries SAC (Special Area of Conservation). Burry Inlet received significant conservation attention back in the 1970s, when the UK government gave permission for 10,000 Oystercatchers to be shot, to protect cockle stocks. This decision was taken despite the objections of conservationists in the UK and Norway, the latter being the summer home of many of these birds. The fact that cockle numbers continued to fall after the cull was an embarrassment, suggesting that Oystercatcher predation was not the only factor at play. Fast forward another 25 years, to the start of the period considered in this paper …

The cockle population in the Burry Inlet SPA declined from 1997 to 2004, before an abrupt ‘crash’ in stocks between 2004 and 2010 which was linked to increased mortality in older cockles, which are particularly important to commercial shellfishers. There are suggestions that losses were associated with warmer summers and sewage releases in periods of wet weather. While there has been some recovery since that period, stocks of larger cockles are still very low.

As cockles are a major prey species for Oystercatchers, the loss of larger individuals may place significant pressure on their populations, as has also been seen in the Dutch Wadden Sea and in The Wash SPA in the UK. Studies in the latter area, by BTO scientists and using data from the Wash Wader Research Group, linked declines in Oystercatcher survival and numbers to years of low cockle numbers (Atkinson et al. 2003 and Atkinson et al. 2005).

When trying to understand Oystercatcher responses to cockle changes in the Burry Inlet and the wider Carmarthen Bay SAC, Katharine Bowgen and colleagues had two main data-sets available to them. The Wetland Bird Survey, and the Birds of Estuaries Enquiry that preceded it, have provided fifty years of data on winter numbers for the Burry Inlet and other local and regional sites. The relative importance of different areas within Carmarthen Bay and the Burry Inlet were established using Low Tide Counts (see below).

Ringing operations also contributed data on the movements of marked birds, individual body masses that could be used to assess the condition of trapped birds, retrap information from which survival rates can be estimated, and opportunities to assess how many juveniles and immatures there are within samples of caught birds. November estimates of cockle biomass were available for the period from 1993 to 2008 (NRW/CEFAS).

What happened to Burry Inlet Oystercatchers?

Here are some of the key findings. Please see the paper for methods and full results. The study was funded by CCW (now NRW).

Extracting Oystercatchers from the cannon-net, having made a successful catch
  • Adult Oystercatchers were found to be in better body condition than immatures and juveniles. Non-breeders may spend three or more years in sites such as the Burry Inlet before first returning to breeding areas.
  • There was considerable variation in annual body condition indices. Two features stood out. Condition was lower in 2005, the winter following the crash in cockle stocks, and improved in the following year. A similar bounce-back could be seen after a particularly cold winter (2010).
  • From recapture data, apparent survival post-2000 was positively correlated with total cockle biomass. Apparent adult survival dropped from an average of 99.3% (range 98.3-99.9%) to 78.5% (range 68.5-84.3%) during the years following the crash in cockle stocks (2004), before rising back to 99.5% (range 99.0-99.9%).
  • In a previous BTO report to CCW (Niall Burton, Lucy Wright et al, 2010) the survival impacts of the Burry cockle crash appeared higher. This effect was diluted with the addition of eight extra years of data presented in this paper. Shorter snapshots of data do not fully capture changes in survival rates for long-lived wader species.
  • Cannon-netting Oystercatchers is not easy! The lack of consistent annual catching success and biases associated with, for instance, just catching the edge of a flock (where juveniles tend to be concentrated), are probably reflected in the fact that no recruitment patterns could be established.
  • WeBS Core Count data showed a significant long-term decrease in the population of Oystercatchers wintering in the Burry Inlet and a long-term increase in the population in Carmarthen Bay. During the period 1997 to 2017 there was a correlation of the two sets of figures – as Burry numbers declined, Carmarthen went up and vice versa (see figure below). Counts of Oystercatcher in the Burry Inlet were weakly associated with cockle biomass in the estuary. Carmarthen Bay counts were more strongly linked to Burry Inlet cockle biomass, increasing as cockle supplies dropped in the Inlet.

Take-home messages

In the study by Katharine Bowgen and colleagues, an apparently underexploited area within the Carmarthen Bay SAC became a vital resource when food supplies collapsed in the Oystercatchers’ preferred feeding area.

Understanding how birds can (and may need to) respond to changing food resources is important, given ongoing pressures from shell-fishers and the fact that the distribution of invertebrate prey stocks may be affected by climate change. The ongoing cockle decline in the Burry Inlet is of concern to both the fishermen, reliant on the stocks, and to conservation managers monitoring bird populations. This study suggests that Oystercatchers may be able to adapt during periods of stress but only if alternative foraging areas are available in the local vicinity.

The analysis of long-term datasets allows more accurate understanding of incidents such as the cockle crash investigated here and improves our abilities to manage their effects on longer-lived species such as waders. Only through long-term monitoring is it possible to fully understand the consequences of major changes in species’ resources and how individuals might adapt to cope with their impacts.

Population level effects

The following three WaderTales blogs contain information about how wintering conditions in particular study areas can affect wader populations.

Sanderling migration

In A place to roost, there is a section about the consequences for local Redshank when Cardiff Bay was permanently flooded, with links to three important papers. Birds that moved elsewhere had difficulty in maintaining their body condition in the winter following removal of their feeding habitat and continued to exhibit lowered survival rates in subsequent years.

In Travel advice for Sanderling,there is clear evidence that poor wintering conditions affect survival rates, the probability of breeding in the first summer and the timing of spring arrival in Greenland. All of these three factors can have population-level effects for the species.

In Gap year for sandpipers, we learn that Semipalmated Sandpipers may not breed every year, depending upon the condition they are in when it is time to migrate.

These three stories are all relevant to the Burry situation. Oystercatchers that were in poor condition at the end of a winter, either because they stayed in the Inlet and had fewer resources or because they moved to new sites, may not have had the resources to migrate to their breeding areas in some years or could have migrated later, either of which may reduce the number of potential breeding attempts within a season (see Time to nest again?).

Conservation implications

Although the analyses presented here were undertaken in order to understand what happened in the Burry Inlet, a site designated in part because of the high Oystercatcher counts, the authors emphasise just how important networks of sites are to wider shorebird conservation issues, especially if there is a rapid change in the quality of a core area. It is not sufficient just to protect the very best sites.

This paper could not have been written without the work of volunteer counters and ringers

With coastal wader populations exhibiting long-term declines globally, understanding how they respond to changes in their prey is important, especially given the potential for warming seas to affect invertebrate populations. In this context, the Burry Inlet study demonstrates the value of long-term WeBS counts and the efforts of local ringers. The contribution of volunteers is warmly acknowledged at the end of the paper.

Resilient protected area network enables species adaptation that mitigates the impact of a crash in food supply. Bowgen, K.M., Wright, L.J., Calbrade, N.A., Coker, D., Dodd, S.G., Hainsworth, I., Howells, R.J., Hughes, D.S., Jenks, P., Murphy, M.D., Sanderson, W.G., Taylor, R.C. and Burton, N.H.K. MEPS. DOI:https://doi.org/10.3354/meps13922


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.