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:
- Wait and hope
- Move elsewhere and never return
- Move elsewhere until conditions improve
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.
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).
- 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.
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.
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.
Have you ever seen a colour-ringed Sanderling and perhaps wondered why it spends the non-breeding* season on a British or Irish beach rather than on one in Portugal, Ghana or even further south? Why fly from Greenland to Namibia, a distance of over 20,000 km, when spending the winter months in the UK or Ireland requires a flight of as little as 3,700 km? Perhaps the chance of survival is greater in other countries or perhaps birds that travel further have a larger lifetime breeding output? A paper by Jeroen Reneerkens and colleagues provides some of the answers.
Sanderling were captured and ringed in breeding sites in northeast Greenland, at staging areas during migration (SW Iceland, N Scotland and the Dutch Wadden Sea) and in the non-breeding season (Scotland, England, Portugal, Mauritania, Ghana and Namibia). The different analyses in these studies used data from 5,863 Sanderling, of which 5,220 were individually colour-ringed.
It’s amazing what colour-ring readers have helped to discover but there is much still to learn about the migration strategies of individual waders.
The map alongside clearly demonstrates the strong link between the Wash and Norway. Other interesting things that have been discovered about Oystercatchers:
In the early days of the WWRG, Grey Plovers occurred in much smaller numbers than they do now. Writing in an article about the first 40 years of the Group, Clive Minton told the story of the first catch of 100, made in 1963, that was celebrated with three bottles of champagne provided by the late Hugh Boyd, delivering on an incentive that he had promised.
The biggest catches of Sanderling are generally in the summer, when the Wash is a meeting point for birds from Greenland and Siberia. July can sometimes see catches of 200 or more birds. Traditionally, a Sanderling catch was the curtain-raiser at the start of Wash Week, an opportunity for the whole team to make one catch before splitting into ‘Terrington’ and ‘Lincolnshire’ teams for the rest of the main summer trip.
The latest population estimates suggest that Great Britain has lost 26,000 wintering Redshank in less than a decade, representing a drop of 20%. Perhaps WWRG data can be used to help to explain these declines? Here are some of the things we know:
Winter Turnstone are birds that will head for Greenland and NE Canada in the spring but recoveries of birds in Finland and other Scandinavian countries indicate a passage of continental birds. African recoveries of WWRG-ringed birds probably include birds from Canada/Greenland and Finland/Scandinavia.
Greenshank: The Group supports a colour-ringing project that was initiated by Pete Potts, in Hampshire. 
The Group still aims to maintain its general ringing programme, so that a representative sample of the key species carry rings. 
Ken Smith, Mike Reed and Barry Trevis have been studying a colour-ringed population of Green Sandpipers in Hertfordshire (an English county just north of London) since 1983. The focal point of their activities is Lemsford Spring, a Hertfordshire & Middlesex Wildlife Trust nature reserve. The site is a set of disused watercress beds that provide plenty of freshwater shrimp (Gammarus pulex) in shallow lagoons fed by freshwater springs. Over the years they have been able to use their colour-ringed birds to study return rates from year to year, behaviour and local habitat use. Using radio-tags they have looked at nocturnal activity patterns and more recently they have been using GPS geolocators to find where their birds go to breed.
The highest numbers of Green Sandpipers in southern England occur in late summer, when many passage birds pass through the country, but a few birds stay for the whole winter. The Lemsford studies show that colour-ringed birds have a high return rate from one winter to the next. These birds are actually in the general area of Lemsford for most of the year, only leaving to breed for two or three months. Upon return, in late summer, they use other local sites such as gravel pits, only switching to the watercress beds in the autumn.
Once established in the Hertfordshire area, individual Green Sandpipers were likely to turn up in subsequent winters, with little difference in return-rates between young birds, ringed in their first winter, and adults. There was considerable variation from year to year, with the return rates varying from 68.7 to 100%; the lowest being between 1986/87 and 1987/88, when another local winter site (Cole Green) became unsuitable. Birds using that site presumably went elsewhere or died. The average apparent annual survival rate was 83.5% (or 84.7% if account is taken of the disappearance of Cole Green). In the only other similar study, in northern Germany, the mean return rate was only 56%, with few birds returning in the year after a severe winter. In the Hertfordshire study, there is a correlation between the overall return rate of colour ringed birds from one winter to the next and the number of nights with air frost in the first winter (r = -0.86, n = 7, P < 0.05). These results suggest that birds suffer increased mortality in severe winters.
From observations at Lemsford, it appears that, once territories have been established, local birds are well aware of their boundaries and neighbours. Should a bird stray into a neighbouring bird’s patch, a low-key call is sufficient warning to avoid a disagreement. Ken reports that the big disputes, with chasing and out-and-out aggression, only occur when the territories are being established or when a new bird tries to feed in an established territory. During extremely cold weather, in the 1980s, there would be lots of extra birds attempting to feed in the spring-fed watercress beds, by trying to squeeze into perceived gaps between territories. Such cold conditions are very rare nowadays.
In recent years, the research team have deployed harness-mounted geolocators to ascertain the movements of a small number of individuals during the course of a full annual cycle. Another WaderTales blog discusses migration and the effects of harnesses on the behaviour of individuals. See 
Over the last fifty years, a range of methods have been used to estimate the annual survival rates of waders/shorebirds. These estimates are based on many hours of patient catching and colour-ring reading, making each one too valuable to discard. How can estimates generated in different ways be combined and what can these estimates tell us about the way in which survival rates vary between species and in different parts of the world?
Return rates: the proportion of marked individuals that are recaptured/resighted in subsequent years. Any bird that changes its location before the next resighting period will reduce the estimate of annual survival for the study population, even though it may still be alive.
As expected, survival rates for larger-bodied species are higher than for smaller species.
Published estimates from the Americas and from smaller shorebirds (Actitis, Calidris and Charadrius spp) tend to be underestimated.
A large number of the world’s shorebird populations are in decline and reduced annual survival rates seem to be a cause for concern in some parts of the world. Although it is impressive that the research team were able to find 295 survival estimates for this paper, I reckon that the data exist to calculate a whole lot more. Each of these extra results might not produce a novel paper but it would be great to see them published somewhere, as brief notes in the journal 
This is an IBIS Open Access Review
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