A Whimbrel’s year

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

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

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

Facing the consequences

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

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

The story so far

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

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

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

Seven years of data

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

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

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

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

Whimbrel YY-LL

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

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

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

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

Results

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

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

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

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

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

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

Over the course of a year

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

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

Iceland’s Whimbrel

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

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

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

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

Here’s the link to the paper:

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


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

When mates behave differently

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

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

Iceland’s Oystercatchers

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

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

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

Fieldwork

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

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

Early nesting attempts may be hampered by spring snowfall

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

Who breeds when?

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

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

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

Reproductive performance

Unusually amongst waders, adult Oystercatchers feed their chicks

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

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

What does this all mean?

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

Hatching brood of three

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

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

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

Long-term studies

Verónica Méndez with one of the marked birds

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

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

The full paper can be found here:

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


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

Iceland’s waders need a strategic forestry plan

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

Pressure on Iceland’s breeding waders

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

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

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

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

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

Assessing the potential impacts of trees

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

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

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

Bird communities change around plantations

Snipe densities are highest close to young forests

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

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

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

The bigger picture

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

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

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

An urgent need for action (and inaction!)

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

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

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

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

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

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

To learn more

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

The paper at the heart of this blog is:

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

Other WaderTales blogs that may be of interest:

Forest edges

Work by Aldís Pálsdóttir (pictured right)

Changing agricultural systems in Iceland (work by Lilja Jóhannesdóttir)


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

Power-lines and breeding waders

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

Breeding waders in Iceland

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

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

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

Power lines

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

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

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

Counting the birds

Aldís counting waders on a transect

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

Results

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

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

Implications of the research

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

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

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

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

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

What are the implications for Iceland’s breeding waders?

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

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

This paper is published as:

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

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

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


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

Whimbrels arrive in Iceland

There is something magical about standing in Eyrarbakki in South Iceland, in spring, watching small flocks of Whimbrel come in over the sea. Thanks to geolocators and satellite tags (about which more later) we know that these amazing waders will have been in the air for around five days, since leaving the west coast of Africa. Looking down at the seaweed-covered rocks on the morning of 27th April 2022, we could pick out small groups of new arrivals. Most were resting but one bird was looking for crabs, just as it had been doing in West Africa just a few days previously. A few birds headed off inland while we were watching, making the distinctive seven-note whistle as they left. It sounded almost like a cry of “made it”!

Incoming Whimbrel: Tómas Gunnarsson

Setting the scene

We had been here before. On 22nd April 2008, I did a live broadcast for the BBC Radio 4 programme World on the Move from this very spot, when I described visible migration to Brett Westwood. On that day, we could see Purple Sandpipers and Turnstone feeding on the tide edge, White Wagtails and Meadow Pipits, newly arrived from Iberia, and small flocks of Golden Plover flying in low over the sea. During the programme, a gaggle of 35 Pink-footed Geese flew in strongly from Britain, as did four Arctic Skuas that had spent winter in the Southern Hemisphere.

On 27th April 2022, the skies were not as busy as they had been fourteen years previously. High pressure over the British Isles and northerly winds across the Atlantic seemed largely to have pressed ‘pause’ on migration from Britain & Ireland. There were newly-arrived White Wagtails, Meadow Pipits and Black-headed Gulls feeding on insects emerging from the banks of rotting seaweed, but the stars were definitely the Whimbrel.

These banks of seaweed will be even more important in May, when there will be hundreds of waders refuelling for the next leg of their journeys, with Sanderling and Dunlin on their way to Greenland and some Knot and Turnstone perhaps flying as far as northeast Canada. When the tide is high enough to wash maggots out of the seaweed, you can sometimes see Red-necked Phalaropes along the tide-line, spinning around and picking food off the surface of the water.

Whimbrel migration

We knew that the Whimbrel were on their way because Global Flyway Network had published a map showing the location of ‘Acuno’, a bird that is wearing a satellite tag that was put on in the Bijagós archipelago of Guinea Bissau. It had been logged west and north of Ireland on the previous evening, already 6000 km and 4 days into its migration. Doing the sums, it seems unlikely that it could have reached Eyrarbakki by the time that we were there but perhaps it flew past soon after, sending signals back to mission control that would confirm arrival**.

** An hour after I published this blog, Acuno was found to have diverted to the Faeroes. It may breed there – as many Whimbrel do – or if could have run out of fuel and landed there, to put on some extra grammes of fat. We’ll see whether it resumes migration.

** Three weeks later, Acuno flew to Iceland. How much will it have been disadvantaged by giving up on direct flight?

Only a minority of Whimbrel fly straight from Africa to Iceland in spring although almost all fly directly south in the late summer. Most individuals spend late April and early May in Ireland, the UK or on the west coast of mainland Europe. Ireland is by far the most important staging area. The individuals on Eyrarbakki beach may have been tired but there could be advantages to being an early bird. See Time to nest again? based on a paper by Morrison et al.

There are several WaderTales blogs about Icelandic Whimbrels:

Whimbrels on the move summarised the movements of Icelandic birds, based on reports of ringed and colour-ringed individuals. In the paper upon which the blog was based (Gunnarsson & Guðmundsson) there was a strong suggestion that birds only stop off in Britain & Ireland on the way north. Geolocator-based research by Alves et al showed that at least some birds were flying straight from Iceland to West Africa and that these sea-crossings could be very rapid.

Migrations to and from Africa were investigated further in a paper by Camilo Carneiro et al that was summarised in Iceland to Africa, non-stop. More recently, papers by the same team have shown that the most consistent point of the annual migration story is departure from Africa and discussed the links between weather and phenology. These two papers have appeared as the WaderTales blogs – Whimbrel: time to leave and A Rhapsody of Whimbrel.

The latest blog about this research is Winter conditions for Whimbrel, based on a paper that assesses the influence of winter conditions on subsequent breeding performance.

Searching for Black-tailed Godwits

We had seen a flock of eight Whimbrel, earlier in the morning, when we were checking fields for colour-ringed Black-tailed Godwits. The Whimbrel were gliding into land, about five km from the coast, before seeming to melt into a patch of rough grassland, bleached after the winter snows.

We did not pay the Whimbrel much attention as, on the other side of the road, there were Black-tailed Godwits probing for worms in silage fields that were already green, after a few days of warmth and the liberal addition of fertilizer. This is our target species during spring trips to Iceland. We have discovered that the arrival time of individuals is remarkably consistent from year to year, which initially seemed surprising, given that migration appears to be getting earlier. There is more about this in Why is spring migration getting earlier? based on a paper in Proceedings of the Royal Society B.

One of the fascinating things about visiting Iceland is that no two years are the same. 2022 has been a dry, warm spring, with northerly winds potentially delaying migration from Britain and Ireland, as mentioned earlier. Early-arriving Black-tailed Godwits that were wearing colour-rings were birds that winter in Portugal and France and migrate via the Netherlands. There was a period of helpful winds for these early birds that fly further but get to Iceland earlier. This strategy is discussed in the blog Overtaking on Migration, based on a paper in Oikos.

Looking forwards

The short Icelandic summer provides fantastic conditions for these waders to raise their chicks, although there are concerns as to how agricultural development, increased forestry and infrastructures will affect these species in the future. In June, in just two months’ time, adults will cross the Atlantic. Black-tailed Godwits head for the British Isles and the west coast of continental Europe and Whimbrel will return to West Africa. By August, the next generation will be preparing for the journey south and we will be here in future springs to monitor their return to Iceland.


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

Oystercatcher Migration: the Dad Effect

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

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

The story so far

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

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

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

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

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

Family ties

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

Focusing on chicks

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

Three colour-ringed chicks. Where will they go?

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

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

Results

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

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

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

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

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

What does this mean?

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

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

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

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

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

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

The broader context

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

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

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


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

Why is spring migration getting earlier?

In Iceland, young trend-setting Black-tailed Godwits are changing the timing of spring migration

Flocks of up to 5000 Black-tailed Godwits gather in Alftafjordur in spring: Tómas Gunnarsson

Flocks of up to 5000 Black-tailed Godwits arrive in Alftafjordur (East Iceland) in spring: Tómas Gunnarsson

In recent years, earlier arrival of spring migrants has been widely reported in birds as diverse as swallows and waders but it’s not a universal trend; species such as British Cuckoos and Icelandic Whimbrels have not changed their arrival dates.  Interestingly, many of the species that have not advanced timing tend to be those that are declining.  By thinking about the mechanisms that enable some species to take advantage of earlier spring warming it might be possible to explain how timings and population changes may be linked.

Are individual Black-tailed Godwits arriving earlier each spring? (Photo: Nigel Clark)

Are individual Black-tailed Godwits arriving earlier each spring? (Photo: Nigel Clark)

The simplest way for spring migration to advance would be for individual birds to change their arrival dates, arriving earlier now (either because they have departed earlier or migrated faster) than they did in previous years.  These changes could be facilitated by changes to weather conditions before, during or after migration.  In general, the arrival of short-distance migrant species has advanced more than long-distance species, which has led to suggestions that individual birds are able to assess conditions on their breeding grounds from afar, and to ‘fine-tune’ arrival accordingly. This could explain why long-distance migrants seem less well able to change their schedules than species which have less far to travel.

Figure 1: Changes in first spring arrival dates of six species of waders in southern Iceland from 1988 to 2009 (reproduced from Gunnarsson & Tómasson 2011)

Figure 1: Changes in first spring arrival dates of six species of waders in southern Iceland from 1988 to 2009 (reproduced from Gunnarsson & Tómasson 2011)

This pattern of advances in arrival dates, and greater advances in short-distance migrants is seen in birds arriving into Iceland each spring (Gunnarsson & Tómasson).   By monitoring the first dates of a range of migratory breeding species to the area around Laugarás, an inland village in southern Iceland, over the period 1988 to 2009, Tómas Gunnarson and Gunnar Tómasson showed that species which spend the winter further south than France showed no change in arrival, whilst those from further north in Europe were returning earlier.  The southern group included the only wader in the Gunnarsson & Tómasson study which uses this migration strategy, Whimbrel (see diagram).

GL-YX on a windy day in western Iceland. He has been seen in eleven years. Despite the vagaries of spring weather, his arrival dates have only been spread over nine days (standard deviation 3.5 days) and have not advanced over the period 2003-2015

GL-YX on a windy day in western Iceland. He has been seen in eleven years. Despite the vagaries of spring weather, his arrival dates have only been spread over nine days (standard deviation 3.5 days) and have not advanced over the period 2003-2015

One of the species that has advanced spring arrival (by about two weeks in the last two decades) is the Black-tailed godwit.  Since 2000, we have been recording arrival dates of individually colour-ringed godwits into coastal Iceland – giving us the opportunity to assess whether individuals have indeed brought forward their time of arrival.  By making regular visits to the same sites we have discovered that the dates when we first come across individuals are remarkably consistent.  Although the arrival of the whole population is spread over a five or six week period, the window in which a specific Black-tailed Godwit appears is generally predictable, whether he or she is a bird that we tend to first see in mid-April or mid-May.  There are annual differences, of course, which appear to be linked to periods of adverse weather during the period of the sea-crossing (Gunnarsson et al 2006), from departure points in The Netherlands, The UK Ireland, France and Portugal, but there is no significant trend.

Dates of spring arrival into Iceland of 54 individually marked black-tailed godwits recorded on arrival in between 4 and 8 years, from 1999 to 2012 (reproduced from Gill et al. 2014). Whether an individual arrives early (left-hand birds) or late (right), the sighting dates for each bird are highly consistent.

Dates of spring arrival into Iceland of 54 individually marked black-tailed godwits recorded on arrival in between 4 and 8 years, from 1999 to 2012 (reproduced from Gill et al. 2014). Whether an individual arrives early (left-hand birds) or late (right), the sighting dates for each bird are highly consistent.

timing pop v indivAlthough the arrival date for the population has been advancing at ~0.8 days per year (Gunnarsson & Tómasson 2011), there has been no trend in individual arrival dates (not significantly different to zero days per year); Gill et al. 2014.

Most of the godwit chicks were ringed by groups of volunteers led by Pete Potts and Ruth Croger (Photo: Tómas Gunnarsson)

Most of the Black-tailed Godwit chicks were ringed by groups of volunteers led by Pete Potts and Ruth Croger (Photo: Tómas Gunnarsson)

If individuals are consistent in arrival but the population is advancing, the advance must presumably result from new birds recruiting into the population being earlier-arrivers than recruits from previous years?  Fortunately, there is a second long-running set of data that’s available to answer this question, in a large part because of the efforts of Pete Potts and Ruth Croger of Farlington Ringing Group.  In the period 1999 to 2014, they organised teams of volunteers to ring Black-tailed Godwit chicks in Iceland, with the support of the Icelandic Natural History Museum.  Significant contributions to the total of over 350 colour-ringed chicks were also made by Tómas Gunnarsson and José Alves, while researching the breeding ecology of Black-tailed Godwits for the Universities of East Anglia and Iceland.

Dates of spring arrival into Iceland of 46 individuals hatched in different years and subsequently recorded on spring arrival (reproduced from Gill et al. 2014)

Dates of spring arrival into Iceland of 46 individuals hatched in different years and subsequently recorded on spring arrival (reproduced from Gill et al. 2014)

Wader chick mortality is quite high and there are further losses in the eighteen-month period between autumn departure from Iceland and the first return trip, eighteen months later, so it was wonderful to have a sufficiently big cohort of marked recruits to look at patterns and trends.  For this study, arrival dates for 46 individuals of known hatch year were available for analysis.  As can be seen from the graph, arrival dates of new recruits have been getting earlier, with birds hatched in the last decade arrive around two weeks earlier than individuals hatched in the 1990s.

There are several reasons why recent recruits may be arriving earlier than in previous years, but the most likely is that this is a knock-on effect of advances in godwit laying dates that have occurred in recent decades.  Icelandic godwits nest earlier in warmer springs, and the frequency of warmer springs has increased.  Early fledging may benefit new recruits, by increasing the time available for them to migrate south, locate a good winter site and be in condition to return early when they recruit into the breeding population (Alves et al. 2013 http://www.esajournals.org/doi/abs/10.1890/12-0737.1  http://dx.doi.org/10.1890/12-0737.1). As their arrival date will be consistent thereafter, the overall timing of arrival of the population will advance.

Will this young Black-tailed Godwit contribute to our understanding of the timing of migration? (Photo: Tómas Gunnarsson)

Will this young Black-tailed Godwit contribute to our understanding of the changing timing of migration? (Photo: Tómas Gunnarsson)

Many other studies of different species in which individuals are tracked during migration are showing similar levels of consistency in individual timing of migration.  What then is causing the variation among species in rates of advance?  Long-distance migrants typically arrive later on the breeding grounds and breed quite soon after arrival, while short-distance migrants can have quite large time gaps between arrival and laying, depending on conditions for breeding. Short-distance migrants therefore have more capacity to advance laying dates (because they are on the breeding grounds waiting for suitable conditions), while long-distance migrants, such as Whimbrels in Iceland, arrive later and so cannot breed earlier even in a warmer year. Advances in spring arrival dates may therefore result from advances in laying dates and associated benefits of early fledging for recruits, and lack of advance in long-distance migrants may be a consequence of arriving late and hence being unable to take advantage of early, warm spring conditions.

In the Icelandic subspecies of Black-tailed Godwit, which is expanding in both number and distribution, it is clear that young recruits to the breeding population are driving the advance in timing of migration.  We only know this because of the long-term programme of chick ringing by volunteers and because we have been able to record the timing of individual birds’ migratory activities over a large number of years.  Funding for this work has been provided by the volunteers themselves, NERC , Icelandic Research Council  and EU TMR.

This blog is based upon research presented in the following open access paper:

Gill, J.A., Alves, J.A., Sutherland, W.J., Appleton, G.F., Potts, P.M. & Gunnarsson, T.G. 2013 Why is timing of bird migration advancing when individuals are not? Proceedings of the Royal Society B. , 281, 20132161 

This work has been taken further in two papers that have been covered in later blogs about warming temperatures and generational change.

Please send reports of colour-ringed Black-tailed Godwits to Jenny Gill (j.gill@uea.ac.uk). She will reply with full details of any birds ringed on the Wash or forward your e-mail to colleagues running other schemes.


 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

Migratory decisions for Icelandic Oystercatchers

New Oystercatcher research in Iceland aims to explain how migration patterns change over time and how individual behaviour may drive population change

Which Oystercatchers will cross the Atlantic and which will stay in Iceland? (Tómas Gunarsson)

Which Oystercatchers will cross the Atlantic and which will stay in Iceland? (Tómas Gunarsson)

Most waders leave Iceland in the autumn, with vast numbers heading for the British Isles, coastal Europe and even Africa.  Amongst the exceptions are hundreds of Oystercatchers which ‘tough it out’ in wintry conditions, thereby saving themselves two long trans-Atlantic flights and potentially ensuring that they are able to make a prompt start to the next breeding season. Why do some Oystercatchers migrate when others don’t?  Is it the same birds each year?  Do resident birds have a competitive advantage when it comes to choosing a territory and raising chicks, and do these chicks follow the same migratory strategy as their parents?  In an era of changing weather patterns, can birds change their migratory preferences and how does this happen?  These are some of the questions that are being tackled by Verónica Méndez, as she monitors the movements of colour-ringed individuals in a joint University of East Anglia (UK) / University of Iceland / University of Aveiro (Portugal) project.

Migration in a changing world

Throughout the world, the distribution, abundance and behaviour of species is changing, partially as a response to climatic change. Given that most bird protection relies on a legislative framework that is site-based, changes to bird distributions are going to make life challenging for conservationists.  Will today’s Special Protection Areas be fit for purpose in 20 years’ time?  As they look to the future, conservationists want to be able to set up a network of sites that can flexibly support migratory species, and this entails understanding why and how migratory patterns are changing and might change in the future.  Icelandic Oystercatchers may well provide an ideal model species through which scientists can learn how migratory decisions can influence individual fitness, population demography and species distribution.

“I’m sure that I started a nest scrape somewhere here”. Spring conditions in 2015 were distinctly wintry: Verónica Méndez

There are likely to be different pressures facing resident and migratory Oystercatchers.  Two birds that breed on adjacent territories might have completely different annual cycles.  The bird that spends the winter in Iceland will not have to migrate but its probability of survival may be reduced by harsher winter conditions, possibly with a more limited set of feeding options and certainly with shorter periods of daylight.  In mid-winter, there is 6hr 40 min between first light and dusk in Reykjavik but 9 hours in Bangor, North Wales.  Come the spring, however, the same bird will be well placed to take advantage if spring comes early, whilst his migratory neighbour will face a hazardous sea-crossing, perhaps being delayed by a fortnight or more if strong northerly winds set in.

Individuals matter

An adult Oystercatcher providing food for a growing chick (Tómas Gunnarsson)

An adult Oystercatcher providing food for a growing chick: Tómas Gunnarsson

Although patterns of change in migratory strategies can be detected by looking at timings of first arrival and counting flocks, these changes can only be explained if we understand what individuals are doing.  It will be interesting to see, for instance, if Icelandic-winterers pair up together and what their offspring subsequently do in their first autumn.  Oystercatchers look after their chicks for longer than most waders so perhaps youngsters will be more likely to adopt the strategy of their parents than other species?

To answer these and many other questions, Verónica and colleagues from the Universities of East Anglia, Iceland and Aveiro have been colour-ringing Oystercatcher adults and chicks on the breeding grounds between the north-west and southern lowlands of Iceland.  Both adults and chicks (when big enough) are ringed with two colour rings on the left tarsus and a flag above another colour ring on the right tarsus (metal ring goes on either tibia and is not part of the scheme).  See update below to learn about a second scheme.

Wintering flocks of Oystercatchers will be checked in Iceland but most of the colour-ringed birds are likely to have flown south.  The team therefore need birdwatchers around the British Isles and along the Atlantic coast of Europe to look out for these marked Oystercatchers and to report them to icelandwader@gmail.com

Catching adult Oystercatchers

The study season for an Oystercatcher fieldworker starts in April, when the first nests start to appear.  Once a full clutch of between 1 and 4 eggs has been produced, both parent birds take turns to incubate for between 3 and 4 weeks.  During this period it is possible to catch the adults by temporarily replacing the eggs with plastic eggs and setting a spring trap or walk-in trap over the nest.  Sometimes the second adult can be caught while the first bird is being ringed, as he or she settles down on the unattended eggs.

Juvenile wearing new colour rings: Verónica Méndez

Juvenile wearing new colour-rings: Verónica Méndez

Catching chicks is harder.  Although they can be metal-ringed at an early age, their legs are too short at this stage for additional colour-rings.  By the time that the youngsters are big enough they can run very quickly – or even fly – providing excellent entertainment for the ‘spotter’ in the research team, as the ‘chaser’ is often outwitted and out-run.

What has been learnt so far?

The first adult Oystercatchers were colour-marked in the summer of 2012 and more were ringed in 2013 and 2014.  In the winter of 2014-15, eight migratory individuals were seen in Ireland, Northern Ireland, Wales, Scotland and Shetland and five resident birds were shown to have stayed in Iceland. Additionally, five birds wearing BTO rings that they acquired in Ireland and Wales have been recruited to the colour-ring project, increasing the sample of known migrants.  By the end of 2015, over 200 adults had been colour-ringed, as well as about 100 new chicks, so there should be plenty of birds to look for this winter.

Breeding adult Oystercatcher: Tómas Gunnarsson

Breeding adult Oystercatcher: Tómas Gunnarsson

A century or more of metal-ringing has shown that Icelandic Oystercatchers migrate mostly to the British Isles and West coast of France, but some have been recovered in Germany, The Netherlands, Spain and Portugal, so please check out any Oystercatcher flocks you see in any of these countries.  Although most observations of colour-ringed Oystercatchers will be made during the winter months, young birds may well be spotted in the summer, as Oystercatchers do not start to breed until they are two or more years old.  If some juveniles winter in Iceland perhaps they might be able to breed at a younger age than their migratory cousins?  That would be yet another way for a change in strategy to not only impact on the fitness of the individual but also on the population demography and the overall migratory strategy of the species.

Oystercatchers are also being ringed with two letter engraved rings, along with two colour-rings: Photo Tómas Gunnarsson

Oystercatchers are also being ringed with two letter engraved rings, along with two colour-rings: Photo Tómas Gunnarsson

An update…

Oystercatchers caught in later catches have been ringed with white, grey or green engraved rings, alongside two colour-rings.  If you come across a bird with a yellow engraved ring then it is probably from a project in Dublin Bay.

When researchers from the two projects got together at the 2015 International Wader Study Group Conference in Iceland they realised that five more Dublin Bay birds could be added to the list of Icelandic breeders that are definite migrants.

Please check flocks of Oystercatchers and report any sightings to icelandwader@gmail.com

Photo: Tómas Gunnarsson

Photo: Tómas Gunnarsson


 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

How volcanic eruptions help waders

Areas of Iceland that have experienced high volcanic ash deposition can accommodate up to three times as many breeding waders.

cropped-2-eldgos.jpg

When Tómas Gunarsson began his PhD at the University of East Anglia on breeding Black-tailed Godwits, supervised by Bill Sutherland and Jenny Gill, little was known about the broad-scale patterns of abundance of waders across Iceland.  As a whole, the country is a haven for breeding waders but what breeds where and why?  In the summer of 2001, Tómas, Jenny and I set off by car to collect data on the distribution and abundance of waders and other bird species within lowland Iceland, continuing this Large Scale Survey during the next two summers.  Having surveyed 764 locations across the lowlands of Iceland, we found a great deal of variation in abundance, some of which was explained by local habitat conditions which we described in Large-scale habitat associations of birds in lowland Iceland: Implications for conservation (Biological Conservation 128 (2006) 265 – 275)

However, there remained a large amount of unexplained difference in wader abundance. Some of this variation appeared to be ‘regional’, as abundances tended to be consistently higher or lower in particular parts of the country, but the cause of these regional differences was not clear.

Fast forward to 2010, when we had the great fortune to be present when Eyjafjallajökull, the famous volcano that halted air traffic across Europe, erupted. As the volcanic ash blew south, disrupting flights and dusting parked cars back in the UK, we thought more about the possible effects of periods of ash deposition within Iceland.  Dust deposition became a hot topic for discussion at the University of Iceland, where Tómas was now Director of the South Iceland Research Centre, especially the effects of ash deposits on soil composition and productivity.  A map of historical dust inputs produced by Ólafur Arnalds, a soil scientist at the Agricultural University of Iceland, showed a pattern that looked very familiar to Tómas.  By collaborating with Ólafur and looking again at the Large Scale Survey data, much of the regional aspect of wader distribution could now be explained.  This work was published as Ecosystem recharge by volcanic dust drives broad-scale variation in bird abundance (Ecology and Evolution 5:12 (2015) 2386-2396)

Survey points for the large scale survey are shown as open circles. Reproduced from Gunnarsson et al 2015.

Survey points for the large scale survey are shown as open circles. Reproduced from Gunnarsson et al 2015.

Dust and birds

It turns out that the regional variation in wader abundance in Iceland is closely associated with volcanic dust deposition, which varies greatly around Iceland because of the active volcanic zone that runs diagonally NE to SW, where the North American and European tectonic plates meet. Areas with high dust input support more waders, probably because of the boost to productivity provided by the high levels of nutrient input. Interestingly, this pattern is most apparent in wetlands (which would be expected as high water levels facilitate nutrient deposition) and is not apparent in agricultural land (probably because more recent fertiliser inputs mask the effects of dust).

Volcanic dust deposition has been linked to primary productivity (vegetation growth) but this is the first study we know of to show links all the way to the top of the food chain.  Iceland may show underlying links between geology and biology more strongly than other countries, where landscapes have been managed more intensively and for much longer than here.

Volcanic dust in Iceland is primarily basaltic and subject to rapid weathering, releasing minerals that are important for ecosystem functions in terrestrial systems.  The fertility of the basaltic volcanogenic dust and rapid deposition rates override the importance of organic matter and clay when it comes to the redistribution of nutrients, through nutrient cycling and buffering of soil and pH.  This can in turn affect invertebrate food resources of birds.

Although volcanoes are the primary producers of dust, distribution is not restricted to active volcanic episodes.  Iceland’s thin soils are readily eroded during windy weather, allowing redistribution of nutrients, and dust trapped in snow and glaciers is released during the spring and summer melt.  As glaciers retreat, the net amount of available ash increases.  Major events such as the eruption of Eyjafjallajökull in 2010 have the potential to significantly add to nutrient levels, although much of the 250,000,000 cubic metres released blew south in this case, with huge consequences for air travel.

Not all of the impacts of dust are positive.  A recent paper has shown that the Eyjafjallajökull eruption reduced the productivity of Whimbrel during the next two years.  See Whimbrels & dust by Borgny Katrínardóttir et al.

Not all species are affected in the same way

Although the long-term effects of dust were clearly present for all birds, when considered together, and for the suite of common birds that were widely recorded during the Large Scale Survey, the clearest patterns appeared in the wader data, as shown below.  In dry areas (middle) and wet areas (right) there are far more waders in regions of the country that have experienced more dust deposition.  The number of individuals tripled between the lowest and highest classes of dust deposition.  Unsurprisingly, there were more waders in wetland areas, as they attract higher numbers of species such as Dunlin, Black-tailed Godwits and Redshank. In farmed areas (left), where slurry and artificial fertilisers are used, the local effects of dust deposition are masked.

The variation in mean (± SE) abundance of ) waders in agricultural habitats, dry habitats and wet habitats in areas of lowland Iceland with differing levels of dust deposition rate (7 = highest deposition rate). Reproduced from Gunnarsson et al 2015.

The variation in mean (± SE) abundance of  waders in agricultural habitats, dry habitats and wet habitats in areas of lowland Iceland with differing levels of dust deposition rate (7 = highest deposition rate). Reproduced from Gunnarsson et al 2015.

At the individual level, five of the seven common wader species all showed a significant response to variation in dust deposition rates (Black-tailed Godwit, Whimbrel, Golden Plover, Redshank & Otstercatcher).  The trends for Snipe and Dunlin were not significant.

What will happen in the future?

Iceland is a country with many active volcanoes and there is always the potential for another eruption that could produce a similar ash plume to that of Eyjafjallajökull in 2010, if basaltic lava erupts beneath a glacier.  Until this happens, smaller eruptions will produce new material in smaller quantities, whilst wind and water will redistribute the dust that has already been deposited. Given that most of the major dust sources are located at glacial margins and in floodplains, and that Icelandic glaciers are retreating due to climate warming, it is likely that volcanic dust inputs will increase in the future, which would have a negative effect on air quality in Iceland, but potentially a positive effect on lowland ecosystem productivity in general. This study shows that dust releases can have large-scale and long-lasting effects across food-webs.

This study was published as: Ecosystem recharge by volcanic dust drives broad-scale variation in bird abundance

Tómas Grétar Gunnarsson, Ólafur Arnalds, Graham Appleton, Verónica Méndez and Jennifer A. Gill  Ecology and Evolution 2015; 5(12): 23862396


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

Introducing WaderTales

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

The choice of topics will reflect personal interests, so there will be plenty about Black-tailed Godwits and the international team of scientists who study their behaviours and life-histories.  I hope that these blogs will be of particular interest to the hundreds of people who contribute their sightings of colour-ringed Black-tailed Godwits to the ever-expanding database of movements.

The first blog appeared on Monday 28 September, telling the story of why areas of Iceland that have been subject to higher deposits of volcanic dust can support up to three times as many waders as those that have received least.

Photo: Tómas Gunnarsson

Photo: Tómas Gunnarsson

On 2 October the second blog outlined new research which aims to explain why some Icelandic Oystercatchers migrate while others stay in Iceland – and how this might impact upon our understanding of how migration patterns change.  As this project unfolds, colour-ring sightings from birdwatchers are once more going to be very important.

By the end of October I hope that there will be a piece on the Black-tailed Godwits of Cley and another on the RSPB’s research into Lapwing predation.

Comments and suggestions will always be welcome.

Graham Appleton

GFA in Iceland