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

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

January to June 2022

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

Welsh Oystercatchers

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

Australian stock-take

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

Chick vocalisation

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

Trans-oceanic migration

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

Hiding in the trees

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

Curlew hunting

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

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

Personal appreciation of Whimbrel

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

Power-line problems

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

Conflict with forestry

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

Blogs from previous years

WaderTales blogs in 2021

WaderTales blogs in 2020

WaderTales blogs in 2019

WaderTales blogs in 2018

WaderTales blogs in 2017

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

Iceland’s waders need a strategic forestry plan

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

Pressure on Iceland’s breeding waders

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

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

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

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

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

Assessing the potential impacts of trees

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

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

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

Bird communities change around plantations

Snipe densities are highest close to young forests

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

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

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

The bigger picture

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

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

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

An urgent need for action (and inaction!)

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

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

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

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

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

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

To learn more

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

The paper at the heart of this blog is:

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

Other WaderTales blogs that may be of interest:

Forest edges

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

Work by Aldís Pálsdóttir (pictured right)

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

Changing agricultural systems in Iceland (work by Lilja Jóhannesdóttir)

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

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

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

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

Who eats African Oystercatcher eggs?

As ornithologists, focused as we are upon birds, we perhaps find it hard to fully acknowledge birds’ eggs as key ‘consumables’ in the food web – a ready source of protein for everything from snakes to deer.

Nest cameras have shown that clutches of ground-nesting waders are taken opportunistically, by grazing sheep for instance, but there are also some species – or individuals – that are specialist egg hunters. See Prickly problems for breeding waders (hedgehogs) and Curlews and foxes in East Anglia (sheep).

In their paper in Wader Study, Itxaso Quintana, Rio Button & Les Underhill describe a single-year study of the predation of African Oystercatcher nests on Robben Island, best known for the prison where Nelson Mandela was held for eighteen years, and for its important seabird colonies. Introductions and eradications on Robben Island have created a far-from-natural food-web that is ever-changing. The situation that Itxaso, Rio and Les found in the 2019/20 breeding season included Kelp Gulls ‘protecting’ African Oystercatcher nests from the attentions of Mole Snakes – something that makes little sense unless set in a historical context, as the authors do in their paper.

What a mess!

Homo sapiens has interfered spectacularly on Robben Island, in the same way as in so many other places, with exploitation of seals, the introduction of an array of large mammal species from the mainland during the apartheid era, and the introduction of rabbits, fallow deer and cats. By 2019/20, the rabbits and almost all of the deer had been removed, encouraging the regeneration of ground cover, and over 90% of the cats had been culled. The remaining threats for African Oystercatcher nests were expected to come mostly from Kelp Gulls, native Mole Snakes and the small number of remaining cats.

Checking out an Oystercatcher’s nest on the shoreline

The 550 African Oystercatchers on Robben Island account for 8% of the species’ population, making this a very important breeding site. This total comprises both nesting pairs and non-breeding birds. African Oystercatchers do not migrate, relying on local shellfish supplies year-round. In 2000, the species was classified as ‘near threatened’ but, thanks to the spread of the invasive Mediterranean Mussel along the South African coast, numbers have recovered. Native shellfish might be in trouble but at least the African Oystercatcher can now be considered to be of ‘least concern’!

The 2019-20 breeding season

Robben island has a coastline which is less than 10 km in length, enabling the authors to monitor 158 nesting attempts of what is thought to have been 133 pairs of African Oystercatchers. A further 300 non-breeding oystercatchers were also present. There were three main study sections, corresponding to the north end of the island (63 nests), the south end of the island (64 nests) and the east side (29 nests). The nest success rates of nests in the north and south were very different, with rates in the east being intermediate.

Below the gull colony in the north of the island, 45 out of 63 African Oystercatcher nests were successful (71%) with one failure associated with Mole Snakes.

Away from the gull colony, in the south of the island, only 14 out of 64 African Oystercatcher nests were successful (22%) with 17 failures associated with Mole Snakes and the reason for most failures unknown.

Breeding Kelp Gulls are spreading across the northern part of Robben Island

There was far more evidence of Mole Snake activity in the south of the island, as can be seen in the map, with individual snakes seemingly ‘patrolling the shoreline just above the spring high tide level, where African Oystercatchers lay their eggs’. In the north, where African Oystercatchers nest on the shoreline immediately below the Kelp Gull colony, snakes were much less conspicuous. You can read more about this in the paper.

Two decades of research

The long-term study of African Oystercatchers on Robben Island started in 2001. Since then, the population has increased fourfold and the number of nests has almost doubled. There have been many changes over this period, as you can read in the paper, but the most significant one for African Oystercatchers is the arrival of Mediterranean Mussels, first noticed in South African waters in 1979 and already colonising the shoreline of Robben Island by 2003. These invasive mussels provide more food than native shellfish and the authors suggest that this has fed through into higher oystercatcher nesting densities.

The beak of an African Penguin is enough to deter Mole Snakes

Over the two decades, there has been a massive change in the populations of potential predators. In the early part of the research period, Mole Snakes were considered relatively unimportant, in terms of predation pressure on African Oystercatchers. Introduced cats, however, then became a major problem, with numbers growing between 2001 and 2005, to such an extent that at least 83% of African Oystercatcher nests were predated in the 2004/05 breeding season. Culls in 2005 and 2006, followed by continued controls, have lowered cat numbers to fewer than ten individuals.

There were no Kelp Gulls breeding on Robben Island until 2000/01, when the first five nests were found. More and more birds now make the short commute to Cape Town, to scavenge, and 2829 gull nests were recorded in 2019/20. Like other large Larus gulls, Kelp Gulls have a reputation as egg thieves. However, on Robben Island, where there are few people disturbing nesting African Oystercatchers, and forcing them to leave their nests, the gulls seem to cause few problems. Instead of being a threat, Kelp Gulls attack Mole Snakes, thereby protecting the eggs of the African Oystercatchers.

For Kelp Gulls, it’s only a short commute to Cape Town, where they scavenge for food

Four other species have benefited from reduced cat numbers. As hoped, numbers of Hartlaub’s Gulls and Swift Terns have both increased, while Mole Snakes have experienced higher survival because the eggs of these two species are available in the austral autumn and winter. African Penguins have benefited too, as a consequence of reduced cat predation, and their sharp beaks can deal with Mole Snakes.

What next?

Hatched Oystercatcher chick and another one on the way

The current situation seems to suit all of the species considered here. Removing introduced herbivores has provided more suitable habitat for snakes, and nesting numbers of key seabirds and African Oystercatchers have all increased since most cats were culled.

Robben Island is not a natural ecological system, however. Major perturbations have happened over decades, with introductions, extirpations, culls and the arrival of invasive species. For the moment. the authors see no reason for Kelp Gulls and/or Mole Snakes to be controlled, in order to support African Oystercatcher numbers or protect important seabird populations.

Cold searching the area above the tide-line to locate nests of African Oystercatchers (Mole Snakes do the same)

One thing that seems almost certain is that the situation will change again: the local Mediterranean Mussel population could collapse, for example due to disease; a pollution incident in the busy sea lanes into the port of Cape Town could suddenly impact birds and/or their food supplies; the grinding effects of climate change, particularly the risk of increased storminess, could slowly upset the equilibrium; diseases such as avian influenza, currently affecting Cape Cormorants, could spread further; or the cat population could explode again. Robben Island looks like a fascinating place in which to study complicated predator-prey interactions; here’s hoping that long-term monitoring will continue.

Paper

The full paper is available in Wader Study, the journal of the International Wader Study Group.

African Oystercatchers on Robben Island, South Africa: The 2019/2020 breeding season in its two decadal context.
Itxaso Quintana, Rio Button & Les G. Underhill. Wader Study.

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

On the beach: breeding shorebirds and visiting tourists

Rising sea levels, stormier weather, coastal development and more people are putting increasing pressures upon shorebirds that nest on beaches. A paper about the breeding waders of Norfolk & Suffolk (UK) coasts illustrates the importance of understanding human behaviour when trying to maintain (or create) space for breeding plovers. This paper will be of particular interest to conservationists trying to support breeding populations of species such as Ringed Plover, Kentish Plover, Piping Plover and Snowy Plover.

The problems of disturbance

Around the globe, nesting plovers are being threatened by human disturbance. Local initiatives to reduce these impacts include the installation of electric fences, recruiting volunteer wardens and changing local bylaws. You can learn more about which interventions work from Conservation Evidence – there’s more about this at the end of this blog.

Panning out, to look at the bigger picture, is it possible to determine where conflicts between breeding birds and tourists are likely to occur, so that one can try to resolve the problems before they start? Can this information help to inform planning decisions?

Oystercatcher with chick

Ringed Plovers and Oystercatchers

The coastline of Norfolk and Suffolk should be an ever-changing environment, dominated by sand and shingle beaches. Coastal defences artificially maintain the barrier between sea and land but sea-level rise is predicted to over-top and destroy sea walls during storm events. How will East Anglia’s beach-nesting waders cope with squeezed beaches at the same time as East Anglia is seeing increasing tourist numbers? In a 2020 paper in Global Ecology & Conservation, Jamie Tratalos and colleagues from the University of East Anglia investigated the distributions of nesting Ringed Plovers and Oystercatchers around the beaches of Norfolk and Suffolk, relating settlement patterns to visitor numbers.

Snettisham beach – sunny weather is not great for the local breeding Ringed Plovers

Ringed Plovers and Oystercatchers breed on sand and shingle beaches that are also attractive to people. They are prone to disturbance, especially by dogs that are allowed to run free, as discussed by Gómez-Serrano (2021). Nests can be trampled, incubation can be interrupted and chicks can be killed. Liley and Sutherland (2006) showed that, over a 9 km stretch of Norfolk coastline, Ringed Plovers bred less successfully when exposed to disturbance by beach visitors, and population declines in this species have been linked to human disturbance (Birds in England by Brown & Grice, 2005). Human recreation has also been shown to be associated with reduced breeding success in Eurasian and other oystercatcher species (Tjørve & Tjørve, 2010), and Ens and Underhill (2014) suggest that increased human use of the coastal zone, combined with increased risk of nest flooding and loss of wetlands, may threaten the conservation of oystercatchers around the world.

Ringed Plovers can have several breeding attempts in the course of a summer

UK breeding populations of Ringed Plover have declined in recent decades, from a conservative estimate of approximately 8400 pairs in 1984 to 4070 in 2007 (Conway et al., 2019) and the species is now red-listed (see the WaderTales blog: Nine red-listed UK waders). Oystercatchers have undergone considerable Europe-wide decline in recent decades and the species has been classified as ‘Near Threatened’ globally (IUCN, 2020).

Counting birds and people

In 2003, when the study at the heart of the Tratalos paper was carried out, East Anglia’s beaches between the Wash and the River Stour held about 3% of the UK’s breeding Ringed Plovers, as well as relatively small numbers of breeding Oystercatchers. As part of a bigger climate change research programme, Tratalos et al were keen to understand what drove the distribution of Ringed Plovers and Oystercatchers, in order to be able to include conservation actions in plans to manage the changing coastline of Norfolk and Suffolk, especially associated the abandonment of outer sea defences. Their research was written up in a 2020 paper in Global Ecology & Conservation.

In the study, the authors examined a 212 km stretch of coastline, mapping all breeding pairs of Ringed Plover and Oystercatcher, as well as the environmental characteristics of beaches. Data on the location of bird territories, and the habitats in which they were found, were collected by Dave Showler in the period between early April and mid-June in 2003. Details of survey methods can be found in the paper.

Map data from Bird Atlas 2007-11 (BTO, BirdWatch Ireland and SOC)

Visitor numbers to different beaches were assessed by filming from a light aircraft, flying at an altitude of 150 metres. 38,634 human visitors were mapped from three flights during sunny weekends in April, June and August, when the tide was at approximately mid phase. There were pronounced peaks in visitor numbers along the coastline, with 19 of the 1003 beach sections experiencing over 10 times the average number of visitors and 231 sections hosting none.

The key findings from surveys and analyses were:

• Of just over one thousand 200m sections of beach surveyed, 183 beach sections contained Ringed Plover territories (266 breeding pairs) and 117 contained Oystercatcher territories (223 pairs).
• There were more occupied territories in less-visited areas, for both species. See table relating the visitor index to occupation of sectors. An index of 0.13 means that visitor numbers were 13% of the mean across all sectors.
• No Oystercatchers were found breeding in sectors where the visitor index was higher than 2.8. No Ringed Plovers were found in sectors where the index was above 5.5.
• Ringed Plovers territories were more common in sections that had dunes at the back of the beach and where the beaches were broader at low tide.
• Oystercatchers appeared to need space above the high-water mark, as well as a broad intertidal area.

The associations between territories and habitat enabled the team to predict the number of pairs of waders that might have been present in areas which were highly impacted by visitors. If visitor numbers were reduced to zero across the whole study area, breeding potential could be hugely increased.

Feeding on the mud at low tide – Ringed Plovers and Oystercatchers need a broad intertidal area

• The authors predict that there would have been an additional 90 beach sections where Ringed Plovers could potentially establish territories, suggesting that tourism and the local use of beaches has already removed 33% of Ringed Plover breeding habitat.
• There were 96 sections where breeding Oystercatchers might have been expected to be found, so they have already lost 45% of potential habitat.

Practical considerations

These results suggest that human activity on beaches influence the location of breeding territories of Ringed Plovers and Oystercatchers, with both species using territories where the number of human visitors was relatively low, when considered both at the scale of the whole Norfolk and Suffolk coast, and locally within areas of this coastline.

In the absence of people, there appear to be clear features of the areas that determine if sectors are used by both wader species for breeding. This makes it possible to predict places where increased access could cause problems so that, ideally, tourism might be encouraged in areas that are less likely to be used by breeding waders. Unfortunately, the beaches that are great for red-listed Ringed Plover – with a back-drop of sand dunes, a sandy beach to walk along at high tide and a gently-shelving intertidal area – are also ones that attract people. This makes it harder to create discrete ‘people zones’ and ‘wader zones’ than might otherwise be the case.

Access points create issues for breeding wader but the effects of most visitors are localised. Unless birdwatching or exercising a dog, the typical tourist will not stray more than 300 metres from a carpark, as indicated in the graphic above. In less-disturbed beach sections, where Oystercatchers set up territories, chicks can hide in upper-beach vegetation until parents indicate that it is safe to come out to be fed.

When Emma Coombes (Global Environmental Change, 2009) asked visitors to Norfolk’s beaches what they were looking for, there was a remarkable consistency in the responses from dog-walkers, sun-bathers and birdwatchers. They all wanted to be on remote flat, sandy beaches, with sand dunes. They would appreciate a car park and toilets too. As soon as such facilities are provided, of course, visitor numbers increase, the remoteness is lost and so are breeding waders, unless fences and wardens are introduced.

Winterton-on-Sea beach is promoted as a tourist location with access to a national nature reserve

Planners have few tools available to them, when it comes to protecting stretches of coastline. All that they can control is development (e.g. new roads, housing and tourist accommodation) and facilities such as car parks and toilets. This paper clearly shows the need to understand the local features that are needed by nesting waders and the importance of documenting current distributions, so that local and national planning authorities have the information they need when planning for the future.

Paper in Global Ecology & Conservation

Vulnerable Ringed Plover chick

Regional models of the influence of human disturbance and habitat quality on the distribution of breeding territories of common ringed plover Charadrius hiaticula and Eurasian oystercatcher Haematopus ostralegus. Jamie A. Tratalos, Andy P. Jones, David A. Showler, Jennifer A. Gill, Ian J. Bateman, Robert Sugden, Andrew R. Watkinson & William J. Sutherland.

Conservation evidence

Before trying a new conservation technique on a local patch, it is worth checking out what has been tried elsewhere. A quick visit to the Conservation Evidence website and a search on ‘beaches’ and ‘bird conservation’ produced a list of 26 actions that have been written up in papers or grey literature. Although many of these interventions are more appropriate to tern conservation than shorebird conservation, a few seem to be particularly relevant to people who are considering how to help breeding Charadrius plovers. Five potential actions are assessed as ‘likely to be beneficial’

• Use signs and access restrictions to reduce disturbance at nest sites
• Physically protect nests with individual exclosures/barriers or provide shelters for chicks of waders
• Protect bird nests using electric fencing
• Physically protect nests from predators using non-electric fencing
• Physically protect nests with individual exclosures/barriers or provide shelters for chicks of ground nesting seabirds

Dogs cannot read ‘no entry’ signs, designed to save space for breeding waders and terns

The Conservation Evidence website aims to make scientific research available to conservation practitioners. Anyone considering any of the interventions listed above can see a quick synopsis of what worked (and what didn’t work) in which circumstances. Anyone who has discovered another successful management technique is urged to write up their study – so that it can be added to the database.

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

Waders on the coast

The UK’s coastline is of international importance because of the numbers of waders that it supports. In winter it accommodates over a third of Europe’s wintering Oystercatcher, Ringed Plover, Bar-tailed Godwit and Knot, as well as an increasing number of Sanderling.

Wintering waders on the UK’s estuaries are counted every month but those on the 17,000 km of open coast are only counted once a decade. There are good reasons for this disparity, given the much higher development pressures on estuaries and the need for regular monitoring of sites that are designated and protected. However, this does mean that we have very little information about wintering Purple Sandpipers, the vast majority of which are not covered by monthly Wetland Bird Surveys (WeBS). Over three-quarters of the UK’s Ringed Plovers are missed too, along with over half of the Sanderling and Turnstones and nearly half of the Curlew.

The last Non-estuarine Waterbird Survey took place during the winter of 2015/16, as discussed in the WaderTales blog NEWS and Oystercatchers. Jenny Gill and I undertook counts on Great Cumbrae and along stretches of the Clyde coast, in Scotland, an area we had also covered for the 2006/07 survey. We were concerned to count only 84 waders in 2015, compared to 206 in 2006. Details are in the table alongside. We hoped that 900 other people, walking along a total of 9000 km of the UK’s coastline, had been more successful!

The paper summarising NEWS results for the whole of the UK and making comparisons with previous surveys in 1997/98 and 2006/07 was not published until 2021. In the intervening period, the counts were included in two papers about wintering populations of waterbirds in Great Britain and Ireland, that were discussed in Do population estimates matter? and Ireland’s wintering waders. This blog draws heavily on a Twitter thread from the Wetland Bird Survey and the BTO’s press release. The new paper is published in Bird Study.

The big picture

In December 2015 and January 2016, NEWS III volunteers walked along amazing, long, white beaches, surveyed rocky headlands and scrambled the lengths of boulder-strewn coves. Not every kilometre of the coast could be visited but the fact that 50% coverage was achieved meant that estimates could be made of the whole coastline of the United Kingdom, together with the Isle of Man and the Channel Isles.

In terms of absolute numbers, Scotland has consistently supported the majority of the population across all non-estuarine waterbird surveys for Oystercatcher, Ringed Plover, Golden Plover, Lapwing, Purple Sandpiper, Bar-tailed Godwit, Curlew, Redshank and Turnstone. Although this is likely to reflect the relative length of the coastline for Scotland (12,714 km) compared to England (2,705 km), Wales (1,185 km) and Northern Ireland (328 km), Purple Sandpiper, Curlew, Redshank and Turnstone still appear to show a bias towards Scotland.

Using the information collected during the survey, BTO scientists were able to extrapolate estimates of the numbers of open-coast waders in the different countries of the UK and its island dependencies (see table below). The results are published in the journal Bird Study and summarised in the table below.

To evaluate the potential importance of the open coast, NEWS estimates for Great Britain in 2015/16 were compared to average population estimates. For eight species, the open coastline accounts for over 20% of the winter population. The figure of 113% for Purple Sandpipers suggests that more birds may have been present on the coasts of the UK in 2015/16 than in an average year or that the population estimate needs to be revisited. There are no Lapwings or Golden Plover in the table below, as there is no recent, reliable estimate of the national wintering population for either species. The Greenshank line is in italics as the sample size is small.

Ten species are considered in detail in the following sections. The maps were downloaded from the BTO website on 20 March 2021 (https://www.bto.org/our-science/projects/ringing/publications/online-ringing-reports). Comparisons are made between results from the Wetland Bird Survey (WeBS) and the Non-estuarine Waterbirds Survey (NEWS).

Oystercatcher

26% use open coasts. 21% NEWS decline since 1997/98. (WeBS decline 22%).

In December 2015, as we walked around the coast of Great Cumbrae in the Firth of Clyde, pairs of Oystercatchers were already staking out their territories, probably not having travelled anywhere since the previous summer or perhaps even in the last twenty years! Wintering flocks that we saw may well have included breeding birds from inland sites in Scotland, from Iceland and from Norway, together with juveniles and non-breeding sub-adults. NEWS III found that densities of coastal Oystercatchers were highest in Wales but that this is the area in which there had been the biggest declines. Breeding numbers have fallen rapidly in Scotland, as you can read in Oystercatchers: from shingle beach to roof-top.

Lapwing and Golden Plover

There was a 68% drop in Lapwing figures between 1997/98 and 2015/16 and a 59% drop in Golden Plover. NEWS and WeBS counts of Lapwing and Golden Plover are difficult to interpret because birds move readily between the coast and inland fields, in response to local conditions such as lying snow and the wetness of fields. This is further complicated in more prolonged freezing conditions, when flocks of Lapwing fly west and south in search of feeding opportunities.

Grey Plover

3% use open coasts. 71% NEWS decline since 1997/98. (WeBS decline 41%).

The Grey Plovers that we see around the coasts of the UK in December and January breed in Siberia. It has been suggested that one of the reasons for the decline in numbers in Britain & Ireland may be related to new generations of youngsters settling in winter locations on the continental side of the North Sea – a strategy that may now work better, given that winters are not as harsh. It is interesting that losses on open coasts, which many would consider sub-optimal habitats, have been more marked than on estuaries. There’s a WaderTales blog about Grey Plovers.

Ringed Plover

82% use open coasts. 21% NEWS decline since 1997/98. (WeBS decline 47%).

Ringed Plovers are red-listed in the UK because of the decline in winter numbers and the importance of these islands of the hiaticula race. In NEWS III, the vast majority of UK birds were found in Scotland (see earlier table) but densities were highest around the coast of England.  Colour-ring studies in Norfolk showed that breeding individuals can adopt a range of migration plans – some marked birds never left the county and others had winter homes as far away as France, Scotland and Ireland. This dispersal is pretty typical of hiaticula race Ringed Plovers that nest in western Europe and southern Scandinavia. Other races travel very long distances (Well-travelled Ringed Plovers).

Curlew

42% use open coasts. 40% NEWS decline since 1997/98. (WeBS decline 26%).

Large numbers of Curlew arrive in the UK in the autumn, with a strong link between Finland and the estuaries of England and Wales. It is estimated that 20% of Europe’s Curlew winter within the British Isles and any change in numbers has significance for a species that is already listed as near-threatened by BirdLife International. The decline in numbers on open coasts has been greater than that seen in estuaries; it has been suggested that this may relate to the breeding origins of birds using different habitats.

Bar-tailed Godwit

15% use open coasts. 33% NEWS decline since 1997/98. (WeBS decline 21%).

Unlike Black-tailed Godwits, which seek out the gloopiest of mud, Bar-tailed Godwits are perfectly at home on sandy shorelines. Wintering birds are of the race lapponica; these breed in Northern Scandinavia, Finland and western Russia (more here). NEWS III tells us that there has been a larger decline in numbers in coastal areas than on estuaries, perhaps related to the relative suitability of the two habitat types.

Turnstone

68% use open coasts. 29% NEWS decline since 1997/98. (WeBS decline 29%).

Almost all of the UK’s wintering Turnstones are thought to be birds that breed in Greenland and Canada. Declines are consistent between NEWS and WeBS. A Northumberland study has shown that, as numbers have dropped, so birds have withdrawn into areas that are less disturbed by people and dogs (See Disturbed Turnstones). About three-quarters of the UK’s open-coast Turnstones are found in Scotland but they are more thinly spread here than in England.

Sanderling

69% use open coasts. 26% NEWS increase since 1997/98. (WeBS increase 8%).

As discussed in Travel advice for Sanderling, the UK is a pretty good place to spend the winter. Whether the same would have been true for previous generations of Sanderling, that were faced with much colder winters, is open to conjecture. Since 1997/98, the densities of Sanderling in Wales have increased by 712%, by 462% in Scotland and by 85% in England. How long will it be until Sanderling flocks successfully over-winter in Iceland?

Dunlin

6% use open coasts. 51% NEWS decline since 1997/98. (WeBS decline 38%).

Three races of Dunlin can be seen in the UK (as you can read in Which wader, when and why?). Wintering Dunlin are birds of the alpina race, arriving in the UK from Siberia, NW Russia, northern Finland and northern Scandinavia in the late summer. Open coasts around the UK are estimated to accommodate fewer than 20,000 Dunlin. To put this into context, there are six estuaries that each hold more than this total during the winter period.

Purple Sandpiper

Almost all on open coasts. 19% NEWS decline since 1997/98. (WeBS decline 34%).

The rocky coasts of the UK are home to Purple Sandpipers from the Arctic, with a suggestion that North Sea coasts south of Aberdeen mainly play host to birds from Spitsbergen and northern Scandinavia, with Greenland and Canadian birds more likely to be found further north and on the Atlantic coast. Coastal numbers have declined by 19%. The Highland Ringing Group has shown that the number of young Purple Sandpipers has been declining on the Moray Firth, suggesting a period of relatively poor breeding success for birds migrating from the northwest.

Redshank

22% use open coasts. 42% NEWS decline since 1997/98. (WeBS decline 21%).

Perhaps surprisingly, few Redshank cross the North Sea to spend the winter in the UK. Winter flocks are largely made up of home-grown birds and migrants from Iceland. The recent decline in Redshank numbers is thought to be a reflection of changing numbers of British and Irish breeders, although there are no monitoring schemes to provide information about Icelandic birds. Since 1997/98, the number of Redshank on open coasts has dropped by 42% but almost all of the losses have occurred in the period since 2007/08 (37% decline between 2007/08 and 2015/16). Redshank is currently amber-listed in the UK, reflecting falling breeding numbers, but ‘promotion’ to the red list cannot be far off. There is a WaderTales blog about the rapid decline in the number of Redshank breeding on salt-marshes: Redshank – the warden of the marshes.

Summary

The Non-estuarine Waterbird Survey 2015/16 revealed that there have been major declines in abundance of four species since NEWS II in 2007/08, only eight years previously: Lapwing (down 57%), Curlew (down 31%), Redshank (down 37%) and Turnstone (down 32%). Lapwing and Curlew are both red-listed in the UK. The only species to increase is Sanderling (up by 79%).

Given the magnitude of the changes revealed in NEWS III, it is unfortunate that this labour-intensive survey can only be carried out every eight to ten years. Ideally, it might be possible to survey at least a sample of sites on an annual basis. It is certainly to be hoped that funding can be found for NEWS IV within the next few years, and that volunteers will once more be prepared to count waterbirds on beautiful, if exposed, stretches of coastline.

The results of NEWS III are published in a paper in Bird Study:

Wader populations on the United Kingdom’s open coast: results of the 2015/16 Non-Estuarine Waterbird Survey (NEWS-III) and a review of population trends. Humphreys, E.M., Austin, G.E., Frost, T.M., Mellan, H.J., Boersch-Supan, P., Burton, N.H.K. and Balmer, D.E.

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

Oystercatcher Migration: the Dad Effect

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

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

The story so far

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

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

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

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

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

Family ties

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

Focusing on chicks

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

Three colour-ringed chicks. Where will they go?

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

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

Results

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

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

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

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

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

What does this mean?

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

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

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

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

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

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

The broader context

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

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

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

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

Where to nest?

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

The perils of ground-nesting

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

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

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

Iceland: a wader factory

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

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

Dragging a light rope across the vegetation to flush nesting birds

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

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

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

First, find your nest

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

This Golden Plover nest was probably predated by an Arctic Skua

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

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

Which nests survive through to hatching?

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

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

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

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

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

In summary

Golden Plover nest set within a homogeneous habitat matrix

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

The paper at the heart of this blog is:

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

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

@grahamfappleton

 

 

Which Icelandic Oystercatchers cross the Atlantic?

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

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

Iceland’s Oystercatchers

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

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

Colour-marks and isotopes

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

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

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

Oystercatchers moult their chest feathers in late winter

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

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

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

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

Migrant or resident?

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

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

Consistency of migratory tendency

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

Factors influencing individual migratory programmes

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

Assortative mating

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

To migrate or remain in Iceland

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

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

Paper

Please click on the title below to access the paper:

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

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

@grahamfappleton