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.


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.

Remote monitoring of wader habitats

In a 2021 paper in Basic and Applied Ecology, Triin Kaasiku and colleagues use radar remote sensing to assess the continued suitability of large areas of Estonia for breeding waders. Although the focus of the paper is upon the performance of an agri-environment scheme, the results are of broader relevance, as conservation biologists seek to monitor vegetation growth and the encroachment of shrubs and trees, especially associated with a warming climate, afforestation and farmland abandonment.

Ground-nesting waders

Many species of waders breed in open habitats. Dense vegetation can remove feeding and nesting areas, and shrub and tree encroachment provides shelter for predators. Previous WaderTales blogs have discussed the perceived and actual predator effects of patches of woodland on breeding Lapwing populations and the difficulty of removing trees and predators that have been planted in peatland (Trees, predators & breeding waders). At a larger scale, vegetation growth, trees and shrubs are threatening breeding areas in arctic and subarctic regions, as discussed by Tómas Gunnarsson in Losing space for breeding waders. Locally, changes to land management, such as reduced sheep grazing and the abandonment of grouse moors in the British uplands, may have unintended consequences for breeding waders, including Curlew.

In Estonia’s coastal grasslands, horses are part of the grazing regime

Maintaining Estonia’s coastal grassland

Estonia’s farmed, coastal grasslands are semi-natural habitats that are vital for a range of species, particularly breeding waders. They also act as soft barriers that dissipate wave energy along coastlines and capture carbon. Across the European Union, the importance of these areas has been recognised, resulting in agri-environment schemes (AES) that preserve farm incomes while encouraging nature-friendly management methods that are associated with lower yields. 

Redshank nest in a clump of grass

Getting the balance right is tricky – for example, too much grazing can reduce heterogeneity and impacts nesting waders (see Big Foot and the Redshank nest), but too little grazing can mean that coastal marshes become rank, as tall grasses replace low-growing plant communities and scrub and trees start to encroach. In a previous paper – Managing coastal grasslands for an endangered wader species can give positive results only when expanding the area of open landscape – Triin Kaasiku, Riinu Rannap & Tanel Kaart emphasised the importance of open, wet grasslands for breeding Dunlin.

The main objective of coastal grassland management in Estonia, in addition to supporting local farming communities, has been to preserve the breeding habitat of several threatened wader species, particularly Dunlin, Ruff, Black-tailed Godwit and Redshank. Nine of the eleven species that breed in these areas have declined moderately or strongly since 1980. Population changes are believed to have been largely driven by habitat loss and increased predation. These problems are recognised around the whole Baltic coast, which is also experiencing more frequent summer storms and flooding. See Dunlin: tales from the Baltic.

Remote sensing

Cute Curlew chick

The Kaasiku team used freely-available European Space Agency Sentinel-1 Synthetic Aperture Radar (SAR) images to assess habitat suitability of coastal grasslands, as breeding sites for waders across Estonia. Although this technique has been used before on cut grasslands, it had not been tested on less uniform grazed areas. The aims of the project were to:

  • Determine the ability of SAR satellite images to classify suitable and unsuitable areas for breeding waders.
  • Compare the habitat suitability for breeding waders under different management schemes, across over 200 km2 of Baltic coastal grassland habitat.
  • Propose methods to improve coastal grassland management.

One of the key parts of the project was to be able to ground truth the SAR data, by visiting sites being managed in different ways (or not at all) in order to collect direct measurements and make observations relating to the suitability of habitat for breeding waders. Details of the methods used are given in the paper.

Research findings

Using SAR radar images, it was possible to correctly identify 88% of suitable habitat and 74% of unsuitable habitat, using a classification boundary that was created using a test set of the data.

Grazing maintains open habitats (left and centre) and restricts reed growth (right)

With renewed confidence, the research team was able to assess whether management prescriptions are actually delivering the desired habitats – for waders and for biodiversity. Their results are encouraging – these agri-environment schemes are broadly working:

  • 43% of Estonia’s coastal grasslands are in a favourable state, in terms of vegetation height.
  • The proportion of suitable habitat is higher (60%) in areas where basic-level conservation subsidies are paid.
  • The highest proportion of suitable habitat (76%) is achieved in areas receiving a top-up conservation subsidy.
  • Long-term grazing ensures better habitat quality: in areas where a basic subsidy has been paid for at least the last five years, 65% of the habitat was classified as suitable.
  • The conclusions were tested by the breeding waders themselves – 98% of the 268 nests located by the team were found in areas classified as suitable.

Although the use of radar remote sensing provides a way of assessing the general suitability of grassland for nesting wader community, it does not provide information on the availability of species-specific microhabitats, finer-scale vegetation characteristics, or soil moisture, all of which are important to breeding waders.

Effectiveness of subsidies

Baltic Dunlin (schinzii) need help

In Estonia, the area of coastal grassland under conservation management has increased from 8,000 to 11,500 ha since the implementation of the AES scheme in 2001. Payments account for 40% of farmers’ incomes. While the payments ensure 60% of the habitat is managed suitably, the remaining 40% does not create habitat for breeding wadersthe main target of the AES for coastal grasslands. The authors suggest that two decades of minimal oversight of the outputs of the scheme has led to sub-optimal delivery. This is unsurprising, given the cost of field visits to farms. Perhaps satellite imagery will help in the future, especially if used five years after the onset of conservation management, when the cumulative effects can be detected.  These assessments will also enable top-up payments to be focused upon areas where 75% of the shoreline is free of high vegetation.

The authors note that the current subsidies are not associated with prescribed stocking rates and suggest that this issue needs to be addressed. Using the right levels of grazing at the right time of year in the right places can make a big difference to breeding waders, as mentioned earlier and discussed in Dunlin – tales from the Baltic and the UK-focused blogs Big Foot and the Redshank nest and Redshank – the ‘warden of the marsh. T

Nest-trampling rates seem low in Estonian coastal grazing habitat, suggesting that there may be scope to increase grazing pressure, especially if cattle are only released onto coastal grasslands in late May.

A broader context

Ruff nest in Estonia

While fencing out predators might be a good way to support breeding waders nesting at high densities in nature reserves or on intensively managed sites, landscape-level support is going to be needed if we are to conserve thinly distributed European species such as Curlew, Dunlin and Golden Plover. Across Europe, prescriptions are in place to adjust grazing levels so as to suit breeding waders but it is difficult to assess the effectiveness of these measures or even to judge whether management is taking place at the right scale and intensity. Triin Kaasiku and collegaues have demonstrated that remote sensing can be a cost-effective way to monitor the delivery of conservation measures indirectly, although they point out that this does not replace detailed monitoring of bird communities. This work has broader applicability, as we seek to measure global-scale habitat changes that will affect increasingly beleaguered wader populations, nesting in some of the most remote parts of the globe.

The paper at the heart of this blog is:

Radar remote sensing as a novel tool to assess the performance of an agri-environment scheme in coastal grasslands. Triin Kaasiku, Jaan Praks, Kaidi Jakobson & Riinu Rannap. Basic and Applied Ecology. July 2021.


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

 

Dunlin: tales from the Baltic

Veli-Matti Pakanen, Kari Koivula  and colleagues have been studying Finnish Dunlin for nearly twenty years. These are schinzii birds that breed in coastal grazing meadows around the Baltic Sea. Several papers have been published, as you will see below, some of which are based on information collected using geolocators attached to leg-flags. Dunlin numbers in coastal Finland are dropping quickly, so this research is important to the conservation of the species. In a 2020 paper, Veli-Matti and colleagues ask whether intensive geolocator-based studies are having a negative effect on individual birds in this already-declining study population.

Global Dunlin

Up to ten races of Dunlin have been identified, which together encircle the globe. Most Dunlin spend the non-breeding season in the northern hemisphere and all migrate north in spring. The breeding and wintering ranges of the various races are summarised at the end of this blog. Here, the focus is on Baltic schinzii Dunlin, a small part of a race that largely winters in coastal North Africa and breeds in southern Scandinavia, Northern Europe, the British Isles, Iceland and southern Greenland.

According to Wetlands International, there were between 4.3 and 6.8 million Dunlin worldwide in 2015, with about one million breeding birds within Europe (BirdLife International). Whilst it is acknowledged that numbers are declining, the large range of the species means that it is still considered to be of ‘least concern’.

Different populations are faring differently. In his description of the changing Dunlin distribution in European Breeding Bird Atlas 2, John Calladine points out that there have been major losses across Europe, including Britain & Ireland, and that Baltic populations ‘are now considered threatened’. The Baltic schinzii population was most recently estimated as between 500 and 640 pairs – less than a fifth of the estimate in the 1980s. John highlights gaps in knowledge that research by the team in Finland and other groups elsewhere are helping to fill.

Migration and survival of schinzii

Dunlin that visit the UK are mostly of the schinzii and alpina races, as indicated in this map. There has been a noticable drop in the number of schinzii birds stopping off in July.

In pre-geolocator days, Ole Thorup and colleagues analysed recovery information available for Dunlin breeding around the Baltic, using 40 years of ringing data from Finland, Sweden, Denmark and Germany. At that stage there were only six mid-winter recoveries in N and NW Africa. The analysis emphasised the importance of wintering and stop-over sites in the Baltic, the Wadden Sea, SE and S England, the Atlantic seaboard of France, and the Iberian Peninsula. Paper in Ardea (2009).

Pakanen et al investigated changes in the survival rates of schinzii Dunlin nesting in Denmark, based on ringing and recaptures of a marked population. They found that annual apparent survival rates dropped from 0.817 to 0.650 between 1990 and 2006, equivalent to a doubling of the chance of dying in any given year. Paper in Bird Study (2016). The importance of monitoring survival rates is discussed in the WaderTales blog: Measuring shorebird survival.

Nests in these flat coastal marshes along the Baltic coast are increasingly susceptible to summer flooding

The use of geolocators enabled Veli-Matti Pakanen to add more detail to the migration story in the 2018 paper, Migration strategies of the Baltic dunlin: rapid jump migration in the autumn but slower skipping type spring migration. He and his colleagues showed that autumn migration is faster than spring migration, characterised by fewer stationary periods, shorter total stopping time and faster flight. The Wadden Sea was found to be an important autumn staging area for all of the tagged birds. Some birds stopped once more before reaching Mauritania. On spring migration, more sites were visited on the way north. The important conservation message from the paper is that Baltic Dunlin may be especially vulnerable to rapid environmental changes at their staging and wintering areas. (In Travel advice for Sanderling there is a suggestion that annual survival is relatively low for birds that winter in Mauritania).

In a 2015 paper in Ornis Fennica, Pakanen et al reported on the results of a single-year analysis of survival rates, concluding that there were no strong effects of leg-flagged geolocators on return rates or reproduction in schinzii Dunlin. However, they did suggest that “long term evaluations that capture the full suite of environmental conditions and assess impact on brood care are needed”. This is a question which leads us neatly on to the 2020 paper: Survival probability in a small shorebird decreases with the time an individual carries a tracking device.

Long-term tracking of individuals

To understand the range of pressures that migratory birds face, one needs to know where individuals spend their time, as was demonstrated in Spoon-billed Sandpiper: Track & Trace and Teenage Waders. Waders of a range of species have been wearing geolocators for ten or more years now, either the same devices on birds which have evaded capture, or a series of tags, when data have been downloaded and replacement tags fitted. These long-term surveys are developing our understanding of the repeatability of migratory behaviour and how birds deal with variable weather patterns, but is there a long-term cost to the individual birds that are tasked with finding out this important information?

This Dunlin wore a ring-mounted geolocator for one year, to collect movement data

Effects of tracking devices on survival are generally considered to be small. However, most studies to date have focused on birds that were caught in one breeding season and recaptured in the following season, to retrieve the geolocator and download the data. In their 2020 paper, Veli-Matti Pakanen and colleagues were able to investigate the possible accumulation of negative effects when individuals have carried the tracking devices for longer periods. Survival rates for tagged birds were compared with 338 colour-ringed birds that were followed for all or part of the period 2002 to 2018.

In the summers of 2013 & 2014, fifty-three adult schinzii Dunlin were fitted with leg-flag mounted geolocators, with a mass equivalent to 1.5 – 2.0% of body-mass. Of these birds, 17 tags were retrieved after one year and 9 after two years. Other marked birds could not be caught and carried their tags for longer periods (2 to 4 years). The research team found that Dunlin carrying a geolocator had reduced chances of survival. Their models suggest that annual survival of colour-ringed males was 0.813. For a bird that carried a geolocator for a year, annual survival probability declined to 0.748 and to 0.581 for birds that carried the geolocator for at least 2 years. Their data suggest that the reduction in survival rates was greater for females than males, even though females are larger than males.

Summer flooding of coastal breeding area is becoming an increasing problem, and likely to get worse with sea-level rise and more chaotic weather patterns

In a thorough Discussion, the authors consider reasons why tags on small waders may be reducing survival, either through ongoing stresses, impacting on things such as feeding efficiency and the energy needed during migration, or because the extra burden means that tagged birds find it harder to cope with occasional periods of tough environmental conditions. They comment on the condition of the skin under removed geolocators – something that other researchers night want to look out for.

As anyone studying breeding waders will know, nest-trapping to retrieve tags is not easy, especially if adults lose their clutches when incubation has only just started, due to flooding, predation etc. Birds may end up carrying tags for longer than intended. The authors “recommend that the detrimental effects of tagging may be avoided by developing attachment methods that are automatically released after one year, e.g. biodegradable materials”.

Balancing costs and benefits

The results from the Pakanen study of long-term survival suggest that requiring a small wader to carry a geolocator for several years may have an impact on survival. As in all mark-recapture studies, researchers are urged to assess the costs to the individual when seeking to understand what might be affecting the viability of a population.

Four previous WaderTales blogs have discussed tag effects:

Details of the Dunlin tagging effects study:

Survival probability in a small shorebird decreases with the time an individual carries a tracking device.

Veli-Matti Pakanen, Nelli Rönkä, Thomson Robert Leslie, Donald Blomqvist, Kari Koivula. Journal of Avian Biology (2020): https://doi.org/10.1111/jav.02555

Four other papers relating to this Finnish Dunlin study

Grazing by cattle is an important management tool in coastal meadows. Pakanen et al studied the impact of trampling on artificial nests and concluded that even recommended stocking rates were too high for chick numbers that could deliver a sustainable population. Paper in Biodiversity and Conservation (2011). (Redshank on British estuaries are similarly vulnerable to trampling – see Big Foot and the Redshank Nest).  

In a follow-up paper in 2016, Pakanen et al concluded that Dunlin populations could be sustained in grazed coastal meadows as long as there was no active grazing before 19 June. Meadows with grazing cattle attracted breeding birds but there was insufficient breeding success for sustainability. Paper in Ecology & Evolution (2016).

If these schinzii Dunlin chicks return to breed they are likely to try to nest nearby; something that needs to be considered when considering conservation measures

Dunlin are strongly philopatric, with both male and female chicks recruiting to suitable habitat close to natal sites. In a paper in Ibis (2017), Pakanen et al show that natal dispersal of Dunlin is strongly linked to the size of their natal site and how isolated the site is. They suggest that inbreeding may be avoided by creating a network of suitably sized patches (20–100 ha sites), no more than 20 km apart from each other. These may work as stepping stones for recruiting individuals. These results are corroborated by a 2021 microsatellite study in BMC Ecology and Evolution which shows genetic differentiation and isolation by distance within the Baltic Dunlin population.

Up to ten races

In western Europe we see three races of Dunlin – alpina, arctica and schinzii. Wintering birds are almost exclusively of the alpina race, which head north and east to northern Scandinavia, Russia and Siberia in spring. The other northern race is arctica, a very small Dunlin that breeds in low numbers in NE Greenland and possibly Spitzbergen. Schinzii has a very large breeding range, spanning the Baltic, southern Scandinavia, Northern Europe, the British Isles, Iceland and SE Greenland. There is huge variation in the timing of breeding of schinzii, as birds do not return to breeding sites in SE Greenland until the end of May, at the same time as schinzii Dunlin being studied by Veli-Matti Pakanen and colleagues in Finland will have their first young chicks.

Further east, centralis Dunlin replace alpina. Many of these birds use the Central Asian Flyway. Further east still, we find sakhalina that use the East Asian/Australasian Flyway (EAA). Two other subspecies have been identified breeding within the EAA Flyway, the more southerly kistchinski birds and actites, which breeds furthest south, on the Russian island of Sakhalin, on a similar latitude to the UK.

It is generally accepted that there are three Dunlin subspecies in North America. In autumn, arcticola head west from northern Alaska and NW Canada and follow the EAA Flyway, pacifica fly south along the Pacific coast from SW Alaska and hudsonia migrate from central northern Canada using the Atlantic Americas Flyway.


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.

More Curlew chicks needed

There are three ways to increase the number of Eurasian Curlew in the UK; boost chick production across the breeding range, find ways to ensure more chicks recruit to the breeding population and/or maximise the lifespan of breeding birds. In a paper in Biological Conservation, Aonghais Cook and colleagues show that, while continued protection of wintering sites is really important, there appears to be little scope for conservation action that can further increase annual survival rates. The focus for conservationists has to be on increasing chick productivity and recruitment.

Curlew in Britain and Ireland

The once-common breeding Curlew is becoming harder to find in many areas. We know that productivity is generally low but could reduced annual survival rates also be contributing to the speed of disappearance? Here’s a quick summary of the story so far.

  • The Eurasian Curlew is designated as ‘near threatened’, as discussed in this WaderTales blog.
  • A 2017 paper by Sam Franks and BTO/RSPB colleagues described the main factors associated with the species’ decline in Great Britain. This work is summarised in a WaderTales blog called Curlews can’t wait for a treatment plan.
  • Estimated breeding population declines since 1995 are 69% in Wales, 59% in Scotland and 31% in England (Breeding Bird Survey). This is not as bad as in Ireland, where 96% were lost between the 1980s and 2015-2017. See Ireland’s Curlew Crisis.
  • Huge numbers of Curlew cross the North Sea at the end of the summer, particularly from Finland. Recent population estimates show that British wintering numbers dropped by 14% in just eight years, with an Irish decline of 13% in five years. See two reviews of wader population estimates, based upon waterbirds papers in British Birds and Irish Birds.

Survival of adult Curlews

As discussed in Measuring shorebird survival, a change in adult survival rates can have a huge effect on shorebird populations. If a species’ annual survival rate drops from 90% to 80% then numbers can half in just six years. We have seen these sorts of dramatic declines in populations of waders that travel between Russia/Alaska and Australia. There’s more about this in Wader declines in the shrinking Yellow Sea.

There are several factors that could be affecting survival rates of British-wintering Curlew:

  • Warmer winters might be expected to lead to increased survival rates.
  • The shooting ban, introduced in 1982 in response to declining Curlew numbers, was specifically designed to increase survival rates.
  • As local wintering populations have dropped, competition for resources could have dropped, potentially leading to increased survival rates.

In their paper about survival rates of Curlew in North Wales, published in Bird Study in 2013, Rachel Taylor and Steve Dodd detected an increase in apparent survival from 86.9% to 90.5% that coincided with the cessation of hunting. They also found that mechanical cockle harvesting in the mid-1990s occurred at the same time as a reduction in apparent survival rate of Curlew from 95% to 81%, indicating the potential for management changes to have huge, unintended consequences for wintering Curlew.

Amassing the data

The research team analysed recoveries of Curlew marked as chicks and adults in the breeding and winter periods, in order to estimate annual survival and the proportion of birds available to recruit into the breeding population.

Thanks to the efforts of skilled, volunteer ringers, two large data-sets were available, collected during the period 1970 to 2018:

  • A total of 1293 adult birds, 144 juveniles and 14,277 chicks were ringed in their breeding grounds across Great Britain.
  • During the winter period, 4403 Curlew were caught and ringed in five key sites – the Severn Estuary, the Tees Estuary, the Wash, Traeth Lavan and the Moray Firth (see map). Supplementary colour-ring sightings were available for the Wash, Tees and Severn.

Full details of how data were used and how models were developed are provided in the paper.

How well do Curlew survive?

Analysis of breeding season data for the period 1970 to 2018 indicates that the average annual survival was 89.8% (confidence interval 0.871–0.920) for adults and 32.6% (0.278–0.378) for first-years. The period of steepest population decline, between 1983 to 1991, coincided with lower survival in both age classes. Encouragingly, since 1996, survival has increased to 92.2% (0.886–0.948) for adults and 39.0% (0.304–0.484) for first-years.

The British and Irish wintering population of Curlew is drawn from a vast area, with birds arriving from as far as Russia and lots of birds from Finland. In a recent breeding wader report covering the Fennoscandia region, no overall change in Curlew numbers was detected, with declines in Norway and Sweden balanced by increases in Finland (see Fennoscandian Wader Factory). From 1970 to 2018, survival rates of UK wintering Curlew averaged 88.4% (0.875–0.893), consistent with survival rates of the British breeding population (above).

Survival varied over time and between the five study sites but has been generally greater than 90% in recent years. Increases in survival were recorded on the Severn Estuary and The Wash.

Curlew struggle in winters with large number of frost-days and survival rates drop significantly, both in that winter and over the subsequent year

Survival was lower in winters with a greater number of days of air frost, an effect exacerbated in successive cold winters. Cold weather may have contributed to low survival in the 1980s, a pattern also evident in the analysis of breeding season data.

Resources may limit numbers. The study suggests that, for four out of five sites, survival was lower in years in which the number of Curlew on a site was higher.

Nationally, the research team found no strong evidence that the hunting ban had increased survival rates. However, there appeared to be local effects on The Severn Estuary and on the Wash. It is unfortunate that national bag data are not available to indicate whether Curlew hunting was particularly prevalent in these two estuaries in the period prior to 1982.

What does this all mean for Curlew conservation?

Birdwatchers are helping to monitor annual survival rates by reporting sightings of colour-marked birds

Since 1996, the mean annual adult survival rate of the British Curlew breeding population has been about 92%. Despite this high number, the Breeding Bird Survey tell us that there has been an observed decline in breeding numbers of 3% per year. Demographic modelling suggests that four breeding pairs must be producing an average of only one chick per year between them. This low figure may come as no surprise to Curlew fans.

To achieve sustainability, the authors conclude that the current figure of 0.25 chicks per pair needs to rise to 0.43 chicks per pair. There are estimated to be 58,000 pairs of Curlew in Great Britain (paper in British Birds). Currently they might be producing about 14,500 chicks each year, on average, and they need to produce nearly 25,000 chicks. British Curlew need to fledge 10,000 more youngsters – every year – just to arrest the decline in numbers.

Personal reflections

Currently, annual survival rates of adults are consistently high but a couple of cold winters, changes to shellfish policy, tidal barrage developments, inappropriately-sited wind turbines and unrestricted disturbance could all have serious negative effects. It is important that we continue to protect the UK’s estuaries and the grassland feeding and roost sites that fall outside their boundaries.

Many Curlew spend significant amounts of time on farmland and recreational land that is not protected in the same way as estuarine habitats

There are a couple of gaps in our knowledge about Curlew demography. When do Curlews first breed and what are survival rates during the ‘teenage’ pre-breeding years? A working hypothesis would be that most breed at age two, with possibly some earlier and probably some later. More colour-ringing of chicks will hopefully provide better data on recruitment age and teenage survival rates. This issue was discussed in Teenage waders. In the meantime, perhaps more thought needs to be applied to avoiding disturbance of non-breeding flocks during the summer holiday season?

It is going to be important to understand how quickly juveniles recruit to breeding populations

It almost goes without saying – Curlews need to produce more chicks. Local conservation initiatives, whether by tenant farmers or dukes with vast estates, will help but raising 10,000 more chicks per year will likely require changes in land management policies. Can agri-environment initiatives be refined to deliver more Curlew? How do we integrate tree-planting and upland conservation priorities? Where should wind turbines be sited? And so much more!

Paper

In their summary of their paper, Aonghais Cook et al concluded that “In addition to increasing productivity, effective conservation strategies will need to maintain high levels of survival, which requires an improved understanding of population connectivity and demographic variation throughout the annual cycle.”

The full paper can be read here:

Temperature and density influence survival in a rapidly declining migratory shorebird.

Aonghais Cook, Niall Burton, Stephen Dodd, Simon Foster, Robert Pell, Robin Ward, Lucy Wright & Robert Robinson. Biological Conservation


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.

Grassland management for Stone-curlew

Detailed studies of a small number of Stone-curlews, breeding in Breckland in the east of England, give some clues as to how to provide the right habitat mix for these big-eyed, nocturnal waders. Increasing structural diversity, by ploughing and/or harrowing areas of grassland, can create an attractive network of nesting and foraging sites for breeding and non-breeding adults.

In a 2021 paper in Animal Conservation, Rob Hawkes and colleagues from the University of East Anglia, RSPB and Natural England give us insights into the daily lives of Stone-curlews nesting in the dry grasslands of East Anglia. By fitting five individuals with GPS tags and following their movements they were able to establish which habitats are used at different stages of the breeding season.

The Stone-curlew is the only migratory member of the thick-knee family. England is at the north-western limit of a breeding range that stretches east to the steppes of Kazakhstan, with birds wintering in southern Europe, North Africa, the Arabian Peninsula and the Indian sub-continent. Most English birds spend the winter months in Spain, Portugal, Morocco or Algeria but a small number are known to cross the Sahara. Stone-curlews return to East Anglia in March and April.

Unmodified grassland in the Brecks of East Anglia provides limited feeding opportunities for Stone-curlews
Creating heterogeneity within areas of grassland by ploughing and/or harrowing patches

During the twentieth century, numbers of Stone-curlew across Europe fell significantly, as mechanized farming expanded. The East Anglian population had dropped to fewer than 100 pairs by 1985 and it took huge conservation efforts to increase this to 200 pairs. Breeding birds are now typically to be found in sparsely-vegetated ground, often in spring-sown crops, on dry heathland or in semi-natural grassland areas, including those used for military training. Preferred food items, such as earthworms, soil-surface invertebrates, slugs and snails, are easier to find in areas of bare and broken ground than in thick grass, so grazing is an important part of conservation action on heaths and in grassland areas.

Each of these square treatments covers a hectare

The UK’s migratory Stone-curlew population has received a huge amount of conservation support, on the back of detailed studies of the species’ breeding ecology (Green et al. 2000).  Tremendous efforts have been made to maximise chick production in farmland, with conservation staff and volunteers working with farmers to monitor breeding pairs, so that they can protect nests and chicks during crop-management operations. In the long term, however, such interventions are too labour-intensive to be sustainable. Can equivalent benefits accrue if more Stone-curlews nest successfully in semi-natural grassland, where the costs of conservation subsidies are lower than in intensively farmed arable cropland?

The study that is reported in the 2021 paper in Animal Conservation took place in an extensive area of semi-natural grassland (nearly 40 km2) that is surrounded by a mosaic of arable farmland. The aim was to understand whether ploughing or harrowing patches of grassland can provide suitable foraging areas, and which other habitats are important.    

Tracking Stone-curlews

GPS tag attached to the back of a Stone Curlew

Stone-curlews are predominantly nocturnal feeders so some form of remote tracking device was needed to understand their movements. GPS loggers were fitted to five adult Stone-curlews during the breeding season. Individuals were caught at night using small, beetle-baited, elastic-powered clap-nets or in the daytime using nest-traps. Each individual was fitted with a 5.2 g solar-powered nanoFix Geo PathTrack GPS tag and an external whip antenna. GPS data were downloaded to a remote base station through a radio connection. Tagged birds were visited at least once a week to establish whether they were still nesting, if they had chicks or had finished breeding.

Where did they go?

One, two and three hectare plots of disturbed ground had already been created within Breckland’s grassland and heathland areas, prior to this study, as described in the paper and discussed in the blog Curlews and foxes in East Anglia. It was already clear that these plots were favoured as nesting sites but does disturbed ground also provide additional feeding opportunities for adult birds – including pairs nesting nearby on arable fields?

Using a telescope to point the base station ‘reader’ at a bird wearing a GPS logger

Three male and two female Stone-curlews were tracked for more than nine weeks (67 to 102 days), yielding 510 GPS fixes during nesting and 1371 post-breeding. There were some fixes in the pre-nesting phase and also during the chick-rearing phase but too few to be considered for analysis. Analytical methods are detailed in the paper.

During the nesting period, what was presumed to be the off-duty bird of each pair was found within 1 km of the nest on 90% of fixes. The mean distance from the nest during daytime feeding was about 100 m but, at night, tagged birds travelled five times as far, on average. They travelled furthest when heading for pig-fields and manure heaps.

  • Relative to closed, undisturbed grassland, nesting Stone-curlews were two- to three-times as likely to forage on disturbed-grassland during night and day, but especially during the day.
  • Night and day, ‘sugar beet or maize’ fields were used more than unmodified-grassland but similar to disturbed-grassland.
  • Nocturnally, Stone-curlews were ten-times as likely to use ‘pig fields or manure heaps’, when compared to undisturbed grassland.
  • The furthest distance travelled was just over 4 km, which is further than previously thought.
Stone-curlews will fly a long way to feed in pig fields

In the post-breeding period, tagged birds travelled up to 13 km to forage, with 90% of nocturnal foraging locations found to be within 5 km of day-time roosting sites.

  • Tagged birds were approximately 15-times as likely to use either disturbed-grassland or arable fallows when compared to undisturbed grassland.
  • Use of the category ‘pig fields or manure heaps’ was nearly as strong (factor c. 10), in comparison to undisturbed grassland.
  • Open crops in the categories ‘sugar beet or maize’ and ‘vegetable or root crops’ were also used more than undisturbed-grassland (factor c. 2).

Conservation messages

In England, there has been a long-term focus on trying to maximise Stone-curlew productivity within arable farmland, especially in East Anglia but also in Wessex. As suggested earlier, this is expensive – foregone production requires high subsidies and interventions by conservation staff are time-consuming. A French study, written up in Ibis by Gaget et al., seriously questions the viability of Stone-curlew populations within intensively managed arable farmland, even with conservation support. Given these problems, should conservation efforts focus on supporting and building up grassland populations?

In a previous tracking study, thirty years previously, Green at al. found that short semi-natural grassland provided suitable foraging habitat for Stone-curlews. Much has changed in Breckland in the intervening period, with the collapse of the rabbit population and an increase in the amount of outdoor pig-rearing. As the short swards of rabbit-grazed grassland have disappeared, Stone-curlews seem to have increasingly taken advantage of alternative opportunities offered in pig fields.

Previous attempts to replicate the grazing efforts of rabbits have involved increasing livestock numbers but this study shows that physical ground-disturbance interventions immediately and effectively create alternative foraging habitat. The authors suggest that multiple areas of disturbed-ground, close to the edge of large grassland blocks, can provide a network of nesting and foraging habitats, whilst allowing access to a mixture of feeding opportunities in the surrounding arable farmland. Of course, nobody is suggesting that intensive outdoor pig-rearing is a positive addition to the habitat mix, as it involves nutrient run-off, ammonia leakage into fragile plant-rich heathland, and pelleted feed attracts corvids (which are known to predate eggs).

The effect of creating ploughed or harrowed plots is almost immediate, in terms of prey availability. In more detailed studies of the effectiveness of slightly different ground-disturbance options, the research team found a strong selection preference only one year after the treatments were first implemented. In the longer term, these areas may become increasingly important, if young birds recruit to the local population and seek out these features. The researchers suggest that a bespoke, ground-disturbance agri-environment option might open up new breeding opportunities within semi-natural grasslands that are currently dominated by tall, closed swords. 

In England, conservation of dry grasslands and heathlands has tended to focus on the preservation of charismatic plant communities, an approach that may have been too gentle and conservative for other taxa. Whilst this study demonstrates that adding pockets of disturbed ground appears to benefit Stone-curlews, previous studies, conducted by this research team, showed benefits for Woodlark (Hawkes et al. 2019), Eurasian Curlew (Zielonka et al. 2019, summarised in Curlews and foxes in East Anglia), and rare, scarce or threatened dry-grassland invertebrates (Hawkes et al. 2019). Similar disturbance techniques have been shown to potentially benefit other grassland-breeding waders, such as Mountain Plovers (Augustine & Skagen 2014) and Upland Sandpipers (Sandercock et al. 2015) in North America, and Sociable Lapwings in Kazakhstan (Kamp et al. 2009).

Paper

This is a summary of a 2021 paper in Animal Conservation:

Effects of experimental land management on habitat use by Eurasian Stone-curlews. Robert W Hawkes, Jennifer Smart, Andy Brown, Rhys E Green, Helen Jones & Paul M Dolman.

Each summer, farmers, volunteers and conservation staff work together to monitor and protect nesting Stone-curlews on farmland, grasslands and heaths in eastern and southern England. Thanks to all of these people, the number of pairs of Stone-curlew in England is holding steady, at over 300 pairs. Progress was reviewed at a conference in 2017, as you can read in this layman’s report and this technical report. More guidance for landowners can be found here.


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.

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Cycling for waders

This blog is mostly about Black-tailed Godwits but there’s stuff about cycling too!

If you’re a Black-tailed Godwit, a 2800-mile (4500-km) direct flight from East Anglia to West Africa is estimated to burn 1085 Calories (4500 kJ) of energy (Alves & Lourenço). Fuelled by Cambridgeshire worms, a female godwit that was raised by the Project Godwit head-starting team flew from the Nene Washes to wetlands in south-east Mauritania in just two days. ‘Cornelia’ – as she was named – undertook this marathon journey with no pre-season training. She just took off on 13th August and arrived on the 15th.

To raise money for Project Godwit and for research projects funded by the International Wader Study Group, Jen and Mark Smart cycled from Somerset to East Anglia, on a 600 mile (960 km) journey that links sites that have been visited by head-started Black-tailed Godwit chicks. Each of them burnt 15,000 Calories (62,800 kJ) over the course of eight days, taking in high energy foods as they travel and stopping to feed and rest each night. Unlike Cornelia, Mark and Jen had been training for years.

Quick reminder of head-starting

Black-tailed Godwits breeding in East Anglia face huge challenges, as you can read below. Four years ago, their situation had become so perilous that it was decided that the only way to stop them disappearing completely was to hatch eggs in incubators and raise chicks in captivity. You can read more about head-starting here. Project Godwit is a partnership between RSPB and WWT, with major funding from the EU LIFE Nature Programme, the HSBC 150th Anniversary Fund, Natural England, the National Lottery Heritage Fund through the Back from the Brink Programme, Leica and the Montague-Panton Animal Welfare Trust.

The maps below show the two breeding sites (Ouse and Nene Washes) and all the late-summer passage sites where head-started birds have been seen in England (left), and the international sightings of all godwits ringed in these breeding sites.

International Wader Study Group (IWSG)

The International Wader Study Group brings together everyone who has a passion for waders (shorebirds), the habitats they use and their conservation. Members include research scientists, citizen scientists and conservation practitioners from all around the world. IWSG gives out small grants each year that help to support wader projects around the world. Recent grants have been used to discover the wintering areas of Common Sandpipers, to measure the site-faithfulness of Dotterel and to support projects in Azerbaijan, Albania, Bangladesh & Argentina.

Mark and Jen

Jen Smart has worked for the RSPB for 14 years.  As a scientist, she led RSPB’s research into the conservation problems faced by breeding waders and developed solutions to help these species. See the WaderTales blog Tool-kit for wader conservation. She developed the science programme around Project Godwit and maintains a keen interest in the project. Jen is Chair of the International Wader Study Group.

‘Manea’ arrived at Old Hall Marshes (Kent) with his sister, ‘Lady’, in July 2017

Mark has worked for the RSPB for 26 years and is Senior Site Manager at Berney Marshes, a 600 ha grassland nature reserve with around 300 pairs of breeding waders. See the WaderTales blog Managing water for waders. As well as managing the reserve, Mark works with other land managers across the country to develop and implement ways of improving habitats for breeding waders.

Latest news from Project Godwit

Project Godwit has been trialling the use of head-starting (https://projectgodwit.org.uk/), where young godwits are reared in captivity, safe from predators and potential flooding, and released once fledged. The aim is to boost the number of godwits breeding in England. The cycle route for Mark and Jen links eleven nature reserves in England, managed by a range of conservation organisations, where head-started Black-tailed Godwits have been spotted on migration by local birdwatchers.

Nelson is one of the birds carrying a geolocator but he has not been recaught (yet)

The ride started at WWT Steart Marshes in Somerset; visited by a Black-tailed Godwit named ‘Nelson’ in 2017. Birdwatchers throughout England were put on alert when the first head-started Black-tailed Godwits were released in 2017 but it was a surprise when Nelson headed southwest. Nelson is a star of Project Godwit. He returned to the Ouse Washes in 2018 and paired up with another head-started bird called ‘Lady’. They have met up in each subsequent spring. In February, Nelson spends time on the Tagus Estuary in Portugal but we don’t know whether he is one of the limosa Black-tailed Godwits that winters south of the Sahara.

The map below shows the route taken by Mark and Jen. The original plan was to cycle from Norfolk to the IWSG conference in Germany, which neatly linked the two causes for which they are seeking sponsorship – Project Godwit and the IWSG fund to support wader research. When the conference was rescheduled as an on-line meeting, they decided to join up the godwit dots across England. The 600-mile bike took just over a week.

Jen & Mark’s route links RSPB, WWT, Wildlife Trustand county wildlife sites between Somerset and Cambridgeshire.

The last site to be visited was the Nene Washes where, as mentioned above, the Black-tailed Godwit ‘Cornelia’ returned to breed. Having been raised at Welney, she was released at the Nene Washes on 27 June, 2018, wearing a small geolocator attached to a flag on her lime ring (see earlier picture). She is the only bird for which the RSPB and WWT team have a whole-year migration history. Cornelia was caught on a nest at the Nene Washes in 2019 and her geolocator was removed. In his blog on the Back from the Brink website, Mo Verhoeven shares his excitement when he learned that this young bird had flown directly from the Nene Washes to wetlands in Mauritania in just two days. There is more about Cornelia here.

Conservation challenges

Wetlands are under threat across the globe and it is appropriate that Mark and Jen are raising money for the International Wader Study Group and for Project Godwit. As they tweeted about their travels and talked about Black-tailed Godwits at local press events, at different nature reserves, they revealed some of the conservation challenges that waders face.

The RSPB nature reserve at Titchwell (North Norfolk) is a favourite pit-stop for Project Godwit birds. These three youngsters, all head-started in 2018, visited before flying south.

Project Godwit is not just about head-starting more Black-tailed Godwit chicks. The team is trying to improve the chances for nesting birds out on the Washes, using electric fences and other predation reduction schemes, and through the development of alternative breeding areas that are under less threat of flooding during spring and summer deluges.

Within Britain and when they head south through Europe and into Africa, Black-tailed Godwits are dependent upon a network of sites. Some of them are fully-protected nature reserves, others have been given international recognition as SPAs and Ramsar sites, but there are many other locations that are important but not designated. Sightings of Project Godwit birds and locations downloaded from geolocators will help to identify areas in which birds may be vulnerable to habitat change and new developments.

A new airport that is planned for the Tagus Estuary is a huge threat to limosa Black-tailed Godwits that breed in Western Europe, including the small English population. It’s thought that about half of the Project Godwit birds use the rice fields and mudflats of the Tagus Estuary, as you can read in Black-tailed Godwits are on their way home. As mentioned above, Nelson has been seen in the Tagus Estuary on several occasions (see map alongside). The proposed airport threatens many species of migrant waterbirds (Tagus Estuary: for birds or planes).

An important unknown when trying to conserve our larger wader species is ‘what happens to the teenagers?’. When do species such as Black-tailed Godwits start to breed and what do they do in the period between fledging and breeding? A key part of Project Godwit is to mark chicks in the wild, as well as head-started birds, hopefully answering questions such as ‘what proportion breed in their first year?’ and ‘where do immature birds spend the pre-breeding years’? Perhaps the International wader Study Group will be able to support similar work for other large shorebird species, through its small project grants?

Support for Mark and Jen

This epic sponsored cycle ride is funding work by Project Godwit and the IWSG. It was a great opportunity to thank colour-ring readers who have reported marked birds, to emphasise the importance of protecting networks of sites for migrant waders, and to highlight some of the conservation challenges that lie ahead.

If you would like to help Mark and Jen to support International Wader Study Group Small Projects Grants, please donate here: https://www.waderstudygroup.org/donate/

If you would like to help Mark and Jen to support the RSPB’s contribution to Project Godwit please donate here: https://www.justgiving.com/fundraising/fundsforwaders


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.

Trees, predators and breeding waders

When trees are planted in open habitats that support breeding waders, numbers usually decline pretty quickly. Trees not only directly remove once-occupied habitat, they are also thought to attract predators, by providing somewhere to hide. In their paper in Restoration Ecology, Mark Hancock and colleagues investigate the distribution of thousands of scats of mammalian predators, in order to understand predator activity in landscapes associated with open bogs and forest edge. They were particularly interested in seeing how long it takes for predators to move out of an area when trees are removed and the land once more reverts to blanket bog.

This study took place alongside a project to repair damage to the vast Flow Country blanket bog (northern Scotland), that occurred in the 1980s, when non-native conifer trees were planted in areas of deep peat that had been drained and deep-ploughed. Such forestry practises would nowadays be prohibited.

More trees

The spread of trees can occur naturally, as the northern treeline moves further north or as trees grow higher in mountain regions, or it can be imposed or accelerated when trees are planted in previously open environments. Warmer temperatures create more opportunities for afforestation and politicians seem to be responding to rising CO2 levels by opting for what seems like an easy win – ‘let’s plant more trees’. In the right places, using appropriate native species, woodland can help to capture carbon**, support woodland wildlife and provide multiple other benefits to society. However, if afforestation focuses on open wet landscapes it can potentially threaten ground-nesting wader species such as Dunlin and Curlew.

** link to The value of habitats of conservation importance to climate change mitigation in the UK by Rob Field and colleagues in Biological Conservation.

As Mark Hancock and colleagues indicate in the abstract of their paper, afforestation of formerly open landscapes can potentially influence mammalian predator communities, with impacts on prey species like ground-nesting birds. In Scotland’s Flow Country, a globally important peatland containing many forestry plantations, earlier studies found reduced densities of breeding waders on open bogs where forestry plantations were present within 700 m. See Hancock et al. 2009 and Wilson et al. 2014. A previous WaderTales blog discussed whether apparent avoidance is due to actual or perceived predation risk (See Mastering Lapwing conservation) but whichever it is, adding new woodland to open wetland habitats has the potential to affect sensitive breeding waders.

Fox scat

There have been many studies looking at the effects of trees on breeding waders but the key differences in this case were that researchers monitored how mammal distributions changed as woodland was removed, in an effort to restore biodiversity and valuable blanket-bog habitats. Spoiler alert: it takes several years to reduce predator numbers!

By counting scats of species as diverse as fox and hedgehog the team were able to address three questions:

  • Did scat distributions vary between open bog, forestry plantations, and former plantations being restored as bog (‘restoration’ habitats)?
  • How fast did scat numbers change in restoration habitats?
  • Were scat numbers different in bogs with differing amounts of nearby forestry?

Counting the poo

Forestry transect

The analyses in this paper are based upon surveying 819 km of track verges, which yielded a total of 2806 scat groups (groups of scats that could have come from one animal) from a variety of predators. I smile when days and days of painstaking fieldwork are summarised in a sentence. “We measured summer scat density and size over 14 years, in 26 transects 0.6-4.5 km in length, collecting data during 93, 96 and 79 transect-years in bog, forestry and restoration habitats respectively”. There is no mention of midges either!

The Flow Country is host to a range of predatory mammals. Hedgehog, Wildcat, Red Fox, Badger, Pine Marten, Stoat and Weasel are native species, while introduced species like Feral Ferret and American Mink may be threatening the area. A 10 mm diameter scat could have been produced by one of six or more species – and increasingly it has been shown that genetic methods, which were outside the budget of this study, are needed to properly identify species from a scat. As all of these species prey on the eggs and/or chicks of breeding waders, the study treated them as a ‘guild’ of animals having similar potential effects.

In April each year, fieldworkers – many of them volunteers based at RSPB’s Forsinard Flows reserve – walked along each of the transects, removing all of the scats from the tracks and track-edges. In July, scats and scat groups that had been deposited on the tracks during the breeding season were counted, measured and their positions recorded using GPS (see paper for details).

Nine transects were in open bog habitat, that remained broadly unchanged throughout the study, 8 were in forestry plantations, 8 were in forestry plantations that were cleared and then restored during the study and 1 site was already a restored plot. For ‘restoration’ transects, the number of years since felling was used as a measure of the length of time under restoration management. In these restoration areas, the brash that was left after felling gradually rotted away or became buried by recovering bog vegetation as re-wetting management (e.g. drain blocking) took effect.

Where’s the poo?

Transect through bog

Predator activity in different habitats and over time. For bog and restoration habitats, scat group density was relatively low throughout the study, averaging around 1 to 2 scat groups per km. In forestry transects, scat group densities started at similar values, but rose approximately eight-fold over the study period, as the forests matured. In the final year of the study, scat group densities in forestry averaged around eight groups per km – approximately twice the figures in restoration and six times the figure in bog habitats. There is a suggestion that more mature forests may have suited Pine Martens, in particular: this species was recorded in the heart of the Flow Country for the first time during the study period.

Trees removed – restoration transect

Predator activity once trees are removed.  Scat group density differed significantly between restoration areas of different age classes. In recently-felled sites (1-5 years), densities were about 2.4 groups per km, rising to 4.0 in the middle period (6-10) and falling to 1.3 later (11-14 years). The authors suggest that tree removal may lead to a flush of nutrients, grasses and then small mammals, thereby explaining the increase in scat densities during the middle stage (6-10 years). The paper demonstrates that it takes several years for mammal densities to fall back to ‘natural’ levels, after tree removal.

Pine Martens have moved into the area

Predator activity close to forests. Scat density on open bog transects was significantly affected by the presence of at least 10% forestry cover within 700 m. There was an estimated 2.9 times (95% confidence limits 1.4 to 6.0) higher scat density at bog transects which contained over 10% cover of forestry within 700 m, compared to bog transects with less forestry nearby. Scottish studies of breeding waders have shown that species such as Golden Plover, Dunlin and Curlew avoid areas close to forestry and the paper includes references to several other similar studies elsewhere.

The Conservation picture

As pointed out by Hancock et al in their Discussion, scat densities in forestry reached much higher values than those of open bogs, especially as the plantations matured, implying that afforestation had strongly altered patterns of mammalian predator occurrence in this formerly open landscape. It took ten years of restoration management to drive down scat densities to levels similar to those of open bogs but, as the authors note, peatland restoration is a rapidly developing field and newer techniques may allow faster restoration, with both biodiversity and soil carbon benefits.

These findings have implications way beyond the scope of this study, three examples of which are included below:

Commercial forestry is seen by some as a way of capturing carbon and can provide opportunities to restore our woodlands, especially our native woodlands and their associated biodiversity. However, given the vulnerability of ground-nesting birds to predation, and the potential for afforestation to markedly affect predator communities, care needs to be exercised when considering afforestation of open landscapes.

Warmer climates offer opportunities to add forestry to the mix of land use options in areas in which the growing season used to be considered too short. For instance, there is significant development pressure in Iceland to plant large areas of non-native commercial forestry. Given that the country holds half or more of Europe’s breeding Whimbrel, Dunlin and Golden Plover this is a contentious issue. This AEWA report is important: Possible impact of Icelandic forestry policy on migratory waterbirds.

In Ireland, it has been suggested that a patchy distribution of relatively new forestry plantations may be one of the factors contributing to the drastic decline of Curlew numbers. (Ireland’s Curlew crisis describes a 96% decline in just 30 years). It has been proposed that some of these patches should be removed. The Hancock et al paper shows how long it might take to make a difference – ten years may simply be too long for Ireland’s breeding Curlew.

Read more

The paper that forms the basis of this blog is:

Guild-level responses by mammalian predators to afforestation and subsequent restoration in a formerly treeless peatland landscape by Mark H. Hancock, Daniela Klein and Neil R. Cowie. Published in Restoration Ecology. https://onlinelibrary.wiley.com/doi/abs/10.1111/rec.13167


GFA in Iceland

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

@grahamfappleton

Scotland’s Dotterel: still hanging on

blogpic brooding

Dotterel brooding chicks

Within the UK, the Dotterel now only breeds on plateaux in the highest Scottish mountains, restricted by habitat that is more commonly found in the arctic or arctic-alpine regions. 

As soon as climate change became apparent, the Dotterel turned into a focal species for ornithologists who were interested in how species would be affected by climate heating. Their fate seemed to be sealed; put simply, there is nowhere colder in Britain to which to retreat when faced with changing habitats and/or breeding conditions.

A 2020 paper by Steven Ewing, Alistair Baxter and colleagues explores the potential ways that changing environmental conditions may be driving the Dotterel’s decline.

Life history

Scottish Dotterels don’t actually spend much time in Scotland, with most birds arriving in early May and leaving within three months. The large part of the year is spent in North Africa, and the plains to the northwest of the Atlas Mountains in Morocco seem to be a particularly important wintering grounds for Scottish birds. Migration north and south appears to be direct, with few European reports of ringed birds in spring and autumn. There is some evidence that Dotterel move further south within North Africa as winter progresses (Whitfield et al 1996), perhaps responding to rainfall patterns.

blogpic map

In May, the numbers of males and females on Scottish breeding sites are roughly equal but many females leave their males sitting on a first clutch of eggs and then depart, leading to an observed drop in sex ratios to about 10:1. Females ringed in Scotland have been spotted breeding with Norwegian males later in the same season and this onward movement to areas with later snow-melt may well be a normal pattern. Indeed, many Dotterels seen on passage in May, often on traditionally used fields or mountain tops south of the Scottish Highlands, may loop north, passing through Highland nesting haunts and then heading northeast into Scandinavia.

A species in decline

blogpic Alistair

Alistair Baxter points to a Dotterel nest that’s right next to a path following the line of a ridge

Dotterels in Britain are at the south-western limit of the species’ global range. They breed almost exclusively in arctic-alpine habitats above 750 m, particularly on Racomitrium moss-heaths that are so characteristic of the flatter topped mountains. These habitats are of high conservation concern, with a tapestry of nationally-rare alpine and arctic plant species.

Scottish Dotterel have been well-studied for over eighty years, a process that was started by Desmond Nethersole-Thompson in the 1940s (detailed in his classic monograph The Dotterel, 1973) and has involved the authors of the Global Change Biology paper since 1987. Some of the areas featured in this paper were studied by Nethersole-Thompson.

An earlier WaderTales blog (Dotterel numbers have fallen by 57%) suggested a number of possible reasons for declines – habitat changes, increased predation and increased disturbance in the Scottish Highlands, compounded by issues affecting the wintering population in North Africa. In the 2020 paper, Ewing et al look in more detail at the potential roles of these changes

Climate and habitat change in Scotland’s mountains

Mountains in Britain are subject to a range of environmental drivers of change that may potentially influence Dotterels, but the logistical challenges presented by working in these environments means that there is rarely good data documenting these changes. This study focuses on snow cover and nitrogen deposition.

blogpic change

The amount of snow-cover is important for cold-adapted species of plants and animals; it insulates the ground in winter and slows up warming in spring, thereby creating a relatively stable environment.  Potential consequences of changes in winter snow-lie for alpine birds might include:

  • A longer growing season for plants, with taller vegetation that reduces the suitability of these areas for species that favour shorter swards.
  • Fewer snow patches, around which Dotterel feed, perhaps also leading to a reduction in peak insect abundance that may not match feeding requirements of chicks.

blogpic nestLots of research carried out in the UK shows that nitrogen deposition is an important driver of upland vegetation change.  Higher deposition of nitrogen tends to result in a reduction of alpine specialist plants, including species of mosses that form key breeding habitats for Dotterel.

The earlier WaderTales blog (Dotterel numbers have fallen by 57%) suggested other possible reasons for Dotterel declines on the breeding grounds, including increased predation and increased disturbance in the Scottish Highlands. While these potential drivers of change could not be tested, due to a lack of data, they are considered in the paper’s Discussion.

Study system

The data that lie at the heart of the Global Change Biology paper have been collected over three decades. Two different but complementary data sources were used in the study.  Firstly, Dotterel were counted at between 128 and 198 alpine sites in the UK during three national surveys in 1987-88, 1999 and 2011.  These censuses focused upon suitable breeding habitats, especially Racomitrium heath, with the latter two surveys successfully covering more than 50% of identified breeding areas.  Secondly, between 1987 and 1999, a smaller cohort of alpine sites were surveyed with far greater frequency (between 40-60 times) as part of SNH’s Montane Ecology Project, where the aim was to study the Dotterel’s breeding ecology in far more detail. The 2020 paper contains detailed information about site use and the parameters that were measured/assessed (elevation, slope, area, snow cover, nitrogen deposition, summer temperature etc.)

blogpic surveyEach site visit involved a lot of climbing, so many of the sites were visited only once per season, with more frequent visits to just 15% of the sites. Having accompanied Phil Whitfield (one of the authors) up one mountain, on one day, I have huge respect for the effort that each data-point represents.  Once up on the tops, observers covered the study areas thoroughly, passing within 100 m of every point and scanning frequently. This has been shown to provide a good count of breeding males.

The authors used their data to investigate whether key potential drivers of environmental change in Scottish mountains (snow-lie, elevated summer temperatures and nitrogen deposition) may have contributed to the population decline of Dotterel.  They also consider the role of rainfall on the species’ wintering grounds in North Africa. The key questions they address are:

  1. Is there evidence of an uphill shift in the elevation of the Dotterel’s breeding range during the study period (1987-2014)?
  2. Are changes in the density or site occupancy of breeding male Dotterels associated with the size, connectedness or topographical aspect of alpine sites?
  3. Does spatial variation in atmospheric nitrogen deposition account for variation in density or occupancy of breeding males at alpine sites?
  4. Are patterns of snow cover or late summer temperatures associated with density or occupancy of male Dotterels at alpine breeding sites?
  5. Do densities of breeding male Dotterels on alpine sites vary with conditions on the North African wintering grounds, as reflected by winter rainfall?

blogpic gloaming

What has changed?

The results are presented in two ways. Data from the period of intensive studies, between 1987 and 1999, are used to try to understand factors influencing annual changes in the number of nesting males. Examination of changes between 1987-90 and 2011-14 gave some indication of factors affecting longer-term trends – something that is important to understand when Dotterel can live for at least ten years.

Densities of breeding male Dotterel in mountainous regions of Scotland declined between 1987 and 1999 and, over the longer-term, site occupancy fell from 80% in 1987 to only 36% in 2014. Densities of breeding males declined disproportionately from lower-lying sites, which resulted in the Dotterel’s breeding range retreating uphill at a rate of 25 m per decade.

Geographically isolated sites appear more likely to lose breeding Dotterel. This makes sense; playback studies in Russia have shown that passing flocks of Dotterel respond to calls, suggesting that birds will be attracted to already-occupied locations.

Settlement patterns were linked to snow-cover.  Generally, Dotterels appear to prefer to settle on higher sites, but late-lying snow at higher elevations appears to deprive them of suitable breeding habitat.  Rather than delay nesting, it seems that these birds then choose to move to lower snow-free sites to breed. Long-term changes in snow cover are poorly documented in high-elevation habitats in Scotland, so it is difficult to know whether the substantial declines observed for Dotterel in recent decades reflect systematic changes in snow-lie.

blogpic snow patch

Nitrogen deposition was shown to be negatively associated with densities of males nesting at lower and intermediate elevations.  The primary impact of nitrogen deposition on Dotterel is likely to be via effects on the species’ favoured Racomitrium moss-heaths, with greater nitrogen levels increasing the rate of moss decomposition and favouring accelerated grass growth.  This presumably results in these habitats becoming increasingly unsuitable for breeding Dotterel.

blogpic chick

Will this chick makes it to Morocco? If it does, how will the conditions it experiences in the non-breeding season affect its probability of return to Scotland?

High rainfall in North Africa seems to lead to higher densities of breeding male Dotterel two springs later, suggesting that wintering ground conditions can potentially influence population dynamics of this alpine-breeding bird.  Similar positive impacts of North African rainfall have also been seen in Ring Ouzels that breed in the UK (Beale et al. 2006).

Dotterel inhabit open farmland and sub-desert steppes in North Africa, where seasonal rainfall brings a flush of vegetation growth and insect abundance. Higher winter rainfall may increase prey availability and Dotterel survival rates but that would be reflected in the arrival numbers in the next spring. The lag of an extra year suggests that low rainfall levels may mostly affect young birds, perhaps delaying recruitment of some Dotterels until their second breeding season.

Conclusions

blogpic juvvyPopulation declines and site abandonment by Dotterel in Scotland during the last three decades have largely occurred at lower elevations, fitting with the traditional idea of climate change limiting the available climate space for alpine breeding species. However, this study found relatively limited evidence that the decline in the breeding population is being driven by climatic factors on the breeding grounds.

Snow cover does seem to influence year-to-year variation in the species’ elevational distribution in Scotland, potentially because a smaller population may now be increasingly settling on higher sites that perhaps were previously unavailable, due to extensive snow cover.  There was also some evidence that greater nitrogen deposition reduced breeding densities of Dotterel at low to intermediate elevations, perhaps by decreasing the suitability of Racomitrium moss heath breeding habitats.  It is also possible that there may have been a redistribution of birds, with newer generations moving further north, to more suitable sites in Norway. (There is a WaderTales blog about this sort of Generational Change mechanism in waders, focusing on Black-tailed Godwit).

Given that Dotterels spend so little time in Scotland, a big gap in our understanding is what is happening in Morocco, where adult Scottish Dotterel spend three-quarters of the year and where young birds may also spend their first summer. How are factors such as rainfall and land-use (particularly farming methods) affecting Dotterels? Might changes in these areas affect other species of migrant that leave northern Europe at the end of the breeding season? Perhaps conservation scientists need to head south for the winter to find out?

Read more in the paper

Clinging on to alpine life: investigating factors driving the uphill range contraction and population decline of a mountain breeding bird. Steven R. Ewing, Alistair Baxter, Jeremy D. Wilson, Daniel B. Hayhow, James Gordon, Des B. A. Thompson, D. Philip Whitfield & René Van der Wal. Global Change Biology.

blogpic dewy


GFA in IcelandWaderTales blogs are written by Graham Appleton, 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.

@GrahamFAppleton

 

Where to nest?

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

The perils of ground-nesting

pic hatching whimbrel

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

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

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

pic hidden Redshank

Iceland: a wader factory

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

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

pic rope

Dragging a light rope across the vegetation to flush nesting birds

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

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

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

pic pretty graph

First, find your nest

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

pic skua-ed goldie eggs

This Golden Plover nest was probably predated by an Arctic Skua

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

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

Which nests survive through to hatching?

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

pic fox attack

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

pic snipe nest

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

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

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

In summary

pic goldie nest in habitat

Golden Plover nest set within a homogeneous habitat matrix

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

The paper at the heart of this blog is:

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

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GFA in Iceland

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

@grahamfappleton

 

 

Curlews and foxes in East Anglia

blog chicksWith Curlew populations in free-fall across much of the British Isles, researchers are trying to understand the reasons for poor breeding performance. At the same time, several groups are trialling emergency interventions, such as predator control and predator exclusion, to try to boost the number of fledged chicks. Sharing knowledge is crucial, so it’s great that a 2020 paper by Natalia Zielonka and colleagues in Bird Study adds to our understanding of nest-site selection and the reasons for nesting failures.

Breckland Curlew

When we think of Britain’s breeding Curlew, the traditional image that comes to mind is moorland, where displaying birds deliver their haunting, bubbling call across upland heather moor and sheep pasture. In lowland East Anglia, in the east of England, things are very different. Some Curlew nest on Breckland heaths, which are structurally similar to moorland, but you can also find nests in sugar-beet field, in military training areas and around airfields.

The Eurasian Curlew is now categorised as Near Threatened by IUCN & BirdLife International, due to populations declines (see Is the Curlew really near-threatened?). Figures from the Breeding Bird Survey for the period 1995-2017 show that the situation in England (30% decline) is less bad than Scotland (down 61%) or Wales (down 68%). The species is now too thinly spread to be monitored in Northern Ireland but we know that the breeding population in the Republic of Ireland dropped from 3,300 pairs to just 138 pairs in 30 years (more in Ireland’s Curlew Crisis). In this context, the hot-spot in Breckland (see left-hand map from Bird Atlas 2007-11) is significant, as is the fact that there are four 10-km squares in East Anglia where an increase in density was noted between 1988-91 and 2008-11 (right).

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Problems for Curlew

The main driver of UK Curlew decline is low breeding productivity, attributable to predation and reduced quality of breeding habitats (see review by Franks et al., summarised in Curlews can’t wait for a treatment plan). Research has informed habitat provision and demonstrated that legal predator control (of foxes, crows, stoats and weasels) can increase Curlew breeding success and abundance – but not in all cases.

blog flyingMuch of the research underpinning the above review was conducted in upland areas. What is happening in the flatlands of East Anglia and might any differences explain the apparent resilience – or even growth – of this population? Most lowland Curlew breed on dry grasslands and heathland, where physical ground-disturbance is increasingly advocated as a land management technique for other rare, scarce and threatened species, such as Stone-Curlew and Woodlark. How do these interventions affect breeding Curlew in the same areas?

Study site

Natalia and her colleagues studied Curlew in two extensive grass-heath sites that differed in predator density and management. Across both sites, vegetation structure had been modified, using ground-disturbance plots, as part of a wider multi-taxa experiment, aimed at increasing biodiversity and supporting key species such as Woodlark (paper in Ibis) and Stone Curlew (paper in Biol. Conserv.).  This disturbance involved creating  2 to 4 hectare plots of deep- or shallow-cultivated ground within the wider grassland/heathland environment.

Before the experiment began, and based on previous research elsewhere, it was assumed that Curlew would avoid physically-disturbed areas, given that Curlew have been shown to prefer to nest in rougher habitats with longer grass swards.  Wherever the Curlews decided to nest, it was predicted that nest survival would be higher on the site with lower predator density, that most clutches would be taken at night and that success would decrease through the season.

blog STANTA

The study was carried out in 2017 and 2018 across the Stanford Military Training Area (STANTA) and Brettenham Heath. Both sites contain extensive areas of dry grassland and grass-heath, surrounded by arable farmland and woodland. Generalist predator control on STANTA was largely carried out around Pheasant release pens, with little or none in most of the surrounding, open arable farmland and woodland. In contrast, Brettenham Heath was subject to continuous predator control across the whole site and in surrounding arable farmland (but not woodland). Brettenham Heath is also enclosed by a two-metre high deer fence with a single electric strand set 50 cm above the ground. There is more about the two sites in the paper (link below).

blog Brettenham

Across both sites, 64 experimental ground-disturbance plots were established in early 2015 and subsequently disturbed annually, using a variety of management techniques. Areas on STANTA that might have contained unexploded ordinance could not be disturbed – and were trickier to survey! In both years, nests were located between mid-April and late June, by visiting any area where Curlew had been seen and looking for adults sitting on, or walking back to, a nest.

Evidence of nest success

blog chickTo determine the date and timing of nest failure, temperature sensors were placed under the eggs. Nests were remotely checked every three-to-seven days, to confirm adults were still incubating, and the scrape was visited once a week to record any predation events (e.g. partial clutch predation). From three days before the predicted hatch date, nests were remotely monitored daily to accurately determine their fate. After hatching, the nest site was visited every three-to-five days, to observe adults and chicks from a vehicle, continuing until the chicks fledged or the breeding attempt had failed.

Three types of evidence can help to reveal the outcome of a nesting attempt:

  • Small chicks or alarming adults are seen in the vicinity of an empty nest.
  • Small shell fragments are found in the nest cup, indicating that chicks have hatched.
  • The temperature sensor reveals that adults kept the eggs warm for the whole incubation period.

A sudden drop in nest temperature can reveal the timing of predation and hint at the culprit. Other studies have shown that nocturnal visits are usually by foxes or badgers, or occasionally hedgehogs. In 2018, infra-red cameras, triggered by movement, were used on ten nests.

blog camera

Where to nest?

Unexpectedly, given that breeding Curlew in the uplands are usually associated with areas with longer grass, Breckland pairs were five- to six-times more likely to select disturbed plots than undisturbed grassland. Nearly half of nests were located on disturbed grassland across both years, which only occupied about 8% of the grassland area. Curlew are long-lived and site-faithful, so disturbed plots may have been created within already-established breeding territories. Whilst physical-disturbance interventions are unlikely to bring birds in from the wider landscape, this study shows that nest placement was more likely to occur on disturbed grassland within a pair’s home range. This suggests that local-scale management can influence nest placement within established breeding sites.

Disturbed grassland is characteristically bare and short compared to uncultivated grassland. Curlew may have placed nests in this habitat because it allows greater vigilance and/or because there is a greater abundance of some important prey for chicks than the surrounding grassland (information in paper by Hawkes et al.). Most nests were on shallow-cultivated plots (created with a rotary rotovator), with few on the barer deep-cultivated plots (created with an agricultural plough). There was taller vegetation and more ground cover in the shallow-cultivated plots.

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Nest survival rates                                                                                                                       

Over the course of the two summers, 44 Curlew nests were monitored, of which 32 failed. 29 nests were predated, one was trampled by livestock, one was mown and one pair deserted (and then renested). Mean overall nest survival probability from start of incubation to hatching was 0.70 ± 0.18 SE at Brettenham Heath (where there was intensive predator control and fencing) and 0.16 ± 0.06 at STANTA (where predator control was patchy and the site was open). Overall breeding productivity was 0.16 ± 0.01 SE fledged chicks per nesting attempt. As predicted, daily nest survival rate decreased through the course of the nesting season.

The figure for STANTA represents low nesting survival, especially when compared to other lowland sites in England, and re-nesting following failure appeared infrequent. Productivity here is likely to be substantially below that required to maintain a stable population. Importantly, nest survival across the two sites was not influenced by ground-disturbance, which suggests that this management intervention did not increase nest exposure to predators.

Identifying the predators

D4_sheep_predation_20May

Sheep versus Curlew

Of the ten 2018 nests with nest cameras:

  • Three nests survived through to hatching.
  • Four were predated by fox (one during the day and three at night).
  • One nest was predated by an unknown predator (following camera malfunction).
  • One nest was predated by a sheep (two out of four eggs remained but incubation was not resumed and the other eggs were later taken by a crow).
  • A single-egg, late-season re-nesting attempt was abandoned three days after camera deployment (egg was later taken by a crow)

There was no effect of nest cameras on daily nest survival rate.

G1_Fox_predation_20May

Fox versus Curlew

Parent Curlews removed some of the temperature loggers but there were sufficient data to identify the timing of predation events for 23 nests. Of these, 17 events were during the night (13 nocturnal, four crepuscular) and six during the day. This nocturnal timing of nest predation was consistent with mammalian rather than avian predation, with camera traps and other evidence suggesting that foxes were the main perpetrators.

As expected, from the levels of predator control, nest survival was lower at STANTA than Brettenham Heath. The latter site was both fenced and subject to lethal fox control, delivering a breeding productivity well above that considered necessary for a sustainable population of Curlew. It is possible that a few fenced sites and others with high levels of predator control might be disproportionately responsible for the fact that Curlew appear to be doing better in the Brecks than in other areas.

Conservation implications

blog bare nestThe key finding of this project is that physical ground-disturbance, which is advocated as a conservation measure within lowland dry grassland and grass-heath for many rare, scarce and threatened species, also provides suitable Curlew nesting habitat, with no reduction in nest survival. Implementing ground-disturbance, particularly through shallow-cultivating, in areas with few or no mammalian nest predators, could provide a useful management tool for attracting breeding Curlew to safer areas.

An intervention to help Stone Curlew and Woodlarks was never designed to assist Curlew. Indeed, there was a prediction that Curlew would actively avoid areas that had been rotovated, in order to create bare patches in which the target species could nest and feed. In a rare case of serendipity, experimental research by Rob Hawkes, Paul Dolman and others has delivered a way of encouraging Curlew to nest in relatively small plots (2-4 hectares) around which it may be possible to run an electric fence. One of the big questions “How can we protect Curlew nests when they have such big territories?” might have become easier to answer. If ‘Curlew plots’ can be created within known territories, or even areas that seem good for Curlew, then they can potentially become the focus for protection.

Having spotted that Curlew seemed to be attracted to disturbed areas it is great that Natalia Zielonka was able to study this population, in order better to understand constraints that seem to be restricting productivity. Her research was undertaken as part of her MSc in Applied Ecology and Conservation at the University of East Anglia,

Further reading

blog RARThe paper at the heart of this blog is:

Placement, survival and predator identity of Eurasian Curlew Numenius arquata nests on lowland grass-heath. Natalia B. Zielonka, Robert W. Hawkes, Helen Jones, Robert J. Burnside & Paul M. Dolman.

Bird Study. DOI 10.1080/00063657.2020.1725421


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