Red Knot pay the price for being fussy eaters!

jean-hall_36310664056_oIn a fascinating comparison of weight gained by Red Knot and Ruddy Turnstone during spring migration in Delaware Bay, on America’s east-coast flyway, Anna Tucker and colleagues show that Knot are far more vulnerable to annual variations in their main food supply than more flexible Turnstones, which target the same food if it is available. Given that changing weather patterns, associated with a warming climate, are expected to make resource availability harder to predict, the authors suggest that populations of migrant shorebirds (waders) that rely on a specific resource being available at the right time are likely to be more vulnerable – as has become apparent for Delaware Bay Knot.

Feast on the beach

Spring shorebird migration in Delaware Bay is largely fuelled by the superabundance of the eggs of horseshoe crabs. Some of the numbers associated with crabs, eggs and migratory waders are amazing!

  • Horseshoe crabs have been around for 400 million years.
  • Each spring, hundreds of thousands of horseshoe crabs swim to Delaware Bay, where the females spawn in the sand along the tidelines of Delaware and New Jersey.
  • Historically, horseshoe crabs were collected, ground up and used as fertilizer. They are still harvested for use as bait and to be bled for the pharmaceutical industry.
  • Each female crab can lay 100,000 eggs, burying them in clumps in the sand above the high tide line, over the course of a few nights.
  • Disturbance by other females and wave action brings eggs to the surface, where they become readily available to birds.
  • In a good year, egg densities on beaches can reach 800,000 per square metre, in hot-spots.
  • Over half a million shorebirds still visit Delaware Bay each spring. This number used to include 100,000 Knot but peak counts are now generally much lower, in the region of 25,000.
  • A Knot can eat 25,000 eggs a day and double its weight in two weeks.
  • Horseshoe crab eggs fuel the second stage of a Knot’s 14,000 km spring migration, from Delaware Bay to the Arctic.

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A short season

In Delaware Bay, horseshoe crabs spawn once a year, during the highest tides in May and early June, a period which overlaps with shorebird stopovers. The largest spawning events typically occur on a full-moon high tide, as long as the water temperature is higher than 15°C. Over the course of two decades, spring water temperatures have fluctuated in Delaware Bay; this annual variability in temperature is predicted to increase with climate change.

Prior to the paper by Anna Tucker and colleagues, which is at the heart of this blog, it had been shown that cooler temperatures cause delays in crab spawn, with consequences for migratory shorebirds. Anna’s research aimed to see whether Knot, which specialise on horseshoe crab eggs, are more susceptible to seasonal differences than Turnstone, which are more generalist foragers.

Knot and Turnstone both breed in the Arctic so they have similarly long journeys ahead of them when they leave Delaware Bay. The prediction was that individual Knot would show more variation in the timing and rate of mass gain than individual Turnstone and that mass gain in Knot, but not Turnstone, would be dependent on the timing of horseshoe crab spawning.

Catch, mark, weigh and re-sight

unnamed (1)Northward migration of Knot and Turnstone begins in late April/early May, with most birds arriving in Delaware Bay in mid to late May and staying for between one and two weeks, before continuing north to begin breeding in early June. The optimal breeding window in the Arctic is short and punctual arrival on territory is important if waders are to have a productive nesting season. This means that spring fattening at migration sites can directly affect breeding success. Weight data for both species were collected between 1 May and 5 June over a twenty-two-year period between 1997 and 2018, with horseshoe crab counts being made since 2003. Birds were marked with flags, with resightings in the same year providing an indication of length of stay. Recaptures of marked birds provided information on weight gain, while sightings in subsequent years gave estimates of apparent survival.

Key results from paper by Tucker et al

For both Knot and Turnstone, spring arrival was pretty consistent between years, over the 22-year period. Approximately 95% of individuals had generally arrived by the time of the seventh sampling period (26–28 May).

The proportion of horseshoe crab spawn that occurred by the time of peak shorebird arrival depended upon water temperature (see paper for details), ranging from just under a quarter (2003) to three-quarters (2004), with a median of 58%. Over the period of the study, there was no sign that the mismatch between horseshoe crab spawning and shorebird arrival was getting stronger.

unnamedMasses of most of the Knot and Turnstone caught in Delaware Bay were within the published range for the two species (Knot 125–205 g and Turnstone 84–190 g) with individually-marked Knot doubling their weight during their stay.

The estimated rate of mass gain per day varied between 2.7 g/day (2005) and 15.9 g/day (1997) for Knot and 2.2 g/day (2014) and 5.8 g/day (1998) for Turnstone. Between years, as predicted, there was far more variability in weight gain for Knot than for Turnstone, with the most pronounced difference being between the maximum rate of weight gain for individuals of the two species. The variability in this parameter for Knot was three times that for Turnstone. For both species, there were higher refuelling rates in years when mass gain started later in the season, with no trend in fattening rates, over the twenty-two-year study, for either species.

Not a lot of Knot

An earlier paper described the decline in Knot numbers in Delaware Bay and the probable link to the availability of horseshoe crab eggs. Rapid population decline in red knots: fitness consequences of decreased refuelling rates and late arrival in Delaware Bay. Allan J. Baker et al. Proceedings of the Royal Society B: Biological Sciences (2004)

  • From 1997 to 2002, an increasing proportion of knots failed to reach threshold departure masses of 180–200 g, possibly because of later arrival in the Bay and food shortage caused by over-harvesting of horseshoe crabs.
  • Resightings suggest that heavier birds in catches made between 1997 to 2002, were more likely to be seen again in subsequent years.

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This 2004 paper and subsequent annual counts of crabs and birds drew attention to overfishing problems in Delaware Bay, leading to a stakeholder-led adaptive horseshoe crab management plan, as described in this paper by McGowan et al. Developing the adaptive management plan required bringing together many groups of stakeholders, with very different values and objectives, to develop a strategy that would allow for the regulated harvest of horseshoe crabs, with a strong emphasis on protecting shorebird populations – especially Knot. The Delaware Bay horseshoe crab population now appears to be stable and it seems that shorebird populations using Delaware Bay have stabilized as well, although at much lower numbers than previously recorded in the Bay.

Conservation initiatives have restricted the impacts of the horseshoe crab fishing industry, reduced disturbance of feeding birds and showcased the importance of Delaware Bay to local people and visitors. You can learn more in this 15-minute programme: Feast on the beach http://www.delmnh.org/feast-on-the-beach/

tableAerial counts of Knot and other shorebirds have been taking place in Delaware Bay since 1981. Over this period Knot counts have varied but the big peaks of nearly 100,000 of the 1980s are now replaced by peaks that rarely exceed 25,000. See table taken from totals reported in USF&WS Rufa Red Knot background information and threats assessment. The declines in the Delaware spring passage numbers are similar in size to the drop in numbers seen wintering in Tierra del Fuego (far south of Argentina).

Why have Knot been so badly affected?

As outlined in the earlier papers and predicted at the start of the new study, the timing of pre-migratory fattening and the rate of mass gain were linked to timing of food abundance for specialist Knot but not for generalist Turnstone. Additionally, mass gain was far more variable in Knot, between years, suggesting greater sensitivity to local conditions (see figure). The way that Turnstones gain mass was more consistent across years and was not associated with abundance or availability of horseshoe crab spawn, which aligns with an ability to dig up buried eggs and switch to alternative prey items. The adaptability of Turnstone is discussed in the blog Why do Turnstone eat chips?

gain graph

The Knot is a highly adapted long-distant migrant. Studies elsewhere have shown that individual Knot can reduce the size of the gizzard, intestinal tract and organs, once they have fattened up for migration. Upon reaching Delaware Bay from southern South America, many Knot will have travelled over 6,000 miles.  The authors suggest that having an available supply of high‐quality soft‐shelled horseshoe crab eggs, which can be consumed without increasing gizzard size, may be particularly important to Delaware Bay Knot. Turnstone are thought more likely to migrate in shorter hops, a strategy that could be physiologically less challenging.

Knot refuelling was slower in years when there were more horseshoe crab eggs, suggesting that a slower mass gain is preferable and that there may be a physiological cost of rapid fat accumulation. Preliminary work (Tucker 2019) suggests that annual survival of Knot appears to be lower in years following springs in which individuals gain weight rapidly.

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In cooler years, with later horseshoe crab spawning, mass gain in Knot was faster, suggesting that birds try to “catch up” at the end of the season, to avoid a delayed departure. It would be interesting to know the upper limit of this extremely rapid mass gain, and whether there are costs to piling on the fat. Such costs could be physiological, in terms of flight efficiency, or there could be increased predation risk if vigilance is compromised while feeding more intensively, for instance.

Breeding shorebirds are constrained by a narrow time window for nesting and a fixed single‐clutch size (usually four eggs). This means that they have little flexibility to compensate for unfavourable conditions previously encountered during migration. In a warming world, with more unpredictable weather patterns, mismatches between food supply and the timing of migratory fattening could well occur more frequently, with downstream consequences for productivity and annual survival rates. The authors point out that this could be particularly bad news for populations of shorebirds that have limited capacity to vary timings within their annual cycles.

In conclusion

unnamed (3)Long‐distance migrants rely on predictable resources at stopover sites; even when these linkages are simple and predictable, populations can be vulnerable to change. The Tucker paper suggests that generalist foraging strategies, as used by Turnstone, may dampen the negative effects of phenological mismatch.

The Knot that use Delaware Bay have probably been relying upon horseshoe crab eggs for centuries and, as long as conditions remain predictable, this remains a good strategy. Over the years, there will have been cold springs, with consequent lower productivity and reduced survival, and other springs in which a superabundance of eggs has fuelled a wonderful summer of eggs, chicks and high survival. Overfishing of horseshoe crabs and more unpredictable spawning conditions appear to have tipped the balance and may (at least in part) explain huge declines in the number of Knot that visit Delaware Bay in spring. This population of Knot has been paying the price for being fussy eaters.

Details of paper

Foraging ecology mediates response to ecological mismatch during migratory stopover. Anna M Tucker, Conor P McGowan, Matthew Catalano, Audrey DeRose-Wilson, Robert A Robinson & Jordan Zimmerman. Ecosphere.

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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.

 

Migration blogs on WaderTales

wwrg GKHere’s a selection of twenty WaderTales blogs that celebrate the wonder of shorebird migration. The focus is mainly on the East Atlantic Flyway, because that’s where I learnt about waders, but there’s a blog about the shocking drop in the numbers of waders that use the Yellow Sea migration route and an amazing story about a shorebird that links Egypt with the far east of Russia. 

Multi-species blogs

tweet long

Oystercatcher

Ringed Plover

  • RP geolocatorWell-travelled Ringed Plovers from Chukotka in north-east Russia spend the winter in Somalia, Egypt, the Red Sea & the Persian Gulf. In Chukotka these Ringed Plovers breed alongside Buff-breasted Sandpipers heading for South America, Knot that will migrate to Australia and Spoon-billed Sandpipers that may migrate to Bangladesh. It’s a small world!

Whimbrel

  • Whimbrels on the move summarises a paper about the movements of Icelandic, ringed Whimbrel, using ringing and colour-ringing data.
  • Iceland to Africa, non-stop discusses the speed of migration of Icelandic Whimbrel in spring and autumn.
  • Whimbrel: time to leave summarises a paper about the consistencies and variability of annual migration patterns of individual Whimbrel.
  • In search of Steppe Whimbrel summarises a migration paper about two very special individual Whimbrel. Will this knowledge help to rescue a subspecies?

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Bar-tailed Godwit

DSCN1827Black-tailed Godwit

The individual movements and breeding season behaviour of Icelandic Black-tailed Godwits have been studied for twenty years. They have been supported by thousands of birdwatchers who report colour-ringed birds.

Sanderling

  • Travel advice for Sanderling summarises research to understand the pros & cons of spending the non-breeding season in widely different locations.

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Snipe & Jack Snipe

Common & Spotted Sandpiper

  • Not-so-Common Sandpipers mixes information about migration with a review of Common & Spotted Sandpipers by Phil Holland.

blog2 movementsGreen Sandpiper

  • Green Sandpipers and Geolocators summarises a Ringing & Migration paper about changing behaviour patterns in Green Sandpipers that wore geolocators to study their migratory journeys.

Spotted Redshank

  • Fewer Spotted Redshanks reviews migration patterns and changes in abundance of the species, in a British & Irish context.

Here’s a link to the full list of WaderTales blogs. https://wadertales.wordpress.com/about/

The intention is to add one or two new blogs each month. You can sign up to receive an e-mail notification when a new one is published.

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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.

Travel advice for Sanderling?

blog 9 wint plum ringedHave you ever seen a colour-ringed Sanderling and perhaps wondered why it spends the non-breeding* season on a British or Irish beach rather than on one in Portugal, Ghana or even further south? Why fly from Greenland to Namibia, a distance of over 20,000 km, when spending the winter months in the UK or Ireland requires a flight of as little as 3,700 km? Perhaps the chance of survival is greater in other countries or perhaps birds that travel further have a larger lifetime breeding output? A paper by Jeroen Reneerkens and colleagues provides some of the answers.

*The term non-breeding season (rather than winter season) is used in this blog because Sanderling travelling as far as Namibia experience a southern summer at the same time as UK birds are experiencing a northern winter.

Pros and cons of travelling further?

At the end of the summer, juvenile Sanderling from Greenland start heading south. The first migration might take an individual to Scotland or Namibia, in southern Africa – or anywhere in-between. The circumstances that lead to these initial settlement patterns are unknown but an individual will repeat its first migratory journey every year, with some birds travelling just 7,400 km annually and others travelling over 44,000 km. It has been argued that, for a range of migration strategies to persist, different wintering sites will have balancing pros and cons. This suggests that costs of longer migrations might be matched by benefits gained at the non-breeding destinations. Is this really true?

blog 5 juv Hebrides

Just six weeks after being ringed as a chick, this Sanderling was photographed in Mull (island off the west coast of mainland Scotland)

Using data provided by colour-ring sightings, Jeroen Reneerkens and colleagues assessed three factors that might affect the fitness of individual birds that spend the non-breeding period in different areas.

  • Annual adult survival: If a bird from one non-breeding location is more likely to survive than a bird that spends the non-breeding season somewhere else, it should live longer and potentially have more breeding attempts within a lifetime.
  • Age when a bird makes its first migration northwards: A young individual that flies north in its first spring will potentially have one more breeding opportunity than a bird that remains in the non-breeding area for its first summer.
  • Timing of migration: There is a short breeding window in the High Arctic so a bird that migrates north earlier in the spring may have higher reproductive success, because it will have a higher chance to re-nest if a first clutch is lost. (There is more about this in Time to nest again?)

Studying marked birds

map no arrowsSanderling were captured and ringed in breeding sites in northeast Greenland, at staging areas during migration (SW Iceland, N Scotland and the Dutch Wadden Sea) and in the non-breeding season (Scotland, England, Portugal, Mauritania, Ghana and Namibia). The different analyses in these studies used data from 5,863 Sanderling, of which 5,220 were individually colour-ringed.

Survival rates. By visiting key sites and collating additional reports of ringed birds from hundreds of birdwatchers, the research team were able to estimate the annual survival rates of Sanderling that spent the non-breeding season in England, France, Portugal, Mauritania, Ghana and Namibia. As can be seen in the table, the apparent survival rate in West Africa (Mauritania and Ghana) was much lower than that in Europe or Namibia. A bird with an annual survival rate of 0.75 is 67% more likely to die in any given year than a bird with a survival rate of 0.85. Confidence limits and methodological notes are provided in the paper. There is a WaderTales blog about the importance of measuring survival rates.

sa plus table

Age of first northward migration. The proportion of colour-ringed juveniles that migrated north in the first spring varied significantly, with virtually all Portuguese and English juveniles migrating north but only 35.8 % of those from Ghana and 9.6 % of those from Mauritania. (There were insufficient data to work out figures for Scotland, France and Namibia).

Timing of northward migration. Observations in Iceland provided information on the timing of migration of Sanderling from a range of non-breeding locations. This is the last possible stop-over site on northward migration, before birds migrate to their Arctic breeding sites in Greenland or Ellesmere Island in Canada. Birds from Ghana were observed in Iceland between 5 and 9 days later and those from Mauritania between 10 to 13 days later than the birds from Europe or Namibia. That is a considerable difference, given the short breeding period.

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Flock of summer plumage Sanderling, on migration in Iceland

Summary

The authors asked the question “is there equal fitness throughout the non-breeding range?”, as inferred from the three measured discussed above. The answer seems to be “no”. Sanderling from non-breeding areas in West Africa had lower annual adult survival, delayed first northward migration and later passage through Iceland than birds wintering either further north or south.

map cross Africa

Sanderling travelling north from Namibia do so by crossing the Sahara (generalised route – sample tracks are shown in paper)

Using geolocators**, the team was able to show that birds from Namibia bypassed potential staging sites in West Africa on the way north, flying north across the African continent to Europe, with some birds stopping briefly in the central part of the Mediterranean before spending a longer stop-over in NW Europe, thereby overtaking the Sanderling from West Africa. Namibian individuals used both Mauritania and Ghana as staging areas during southward migration.

** The use of geolocators is discussed in this blog.

The West African sites seem to be relatively poor places in which to spend the non-breeding months of the year. Food availability in spring is likely to be the chief problem. Theunis Piersma and colleagues have shown that the quality and biomass of prey available to shorebirds is lowest close to the equator, resulting in low fuelling rates and low body masses at departure for northward migration in Knot (Piersma et al. 2005).

Sanderling occupy a variety of different and widely-dispersed non-breeding sites between the northern tip of Scotland and the southern tip of the African continent. Here, they experience very different conditions which affect potential, life-time breeding outputs. Sites which appear to be poorer continue to be used, even though there are better options elsewhere, simply because individual birds have no knowledge of other potential areas where they could spend the non-breeding months.

A roll of the dice?

Once a juvenile Sanderling has settled upon a particular migration strategy and a spot in which to spend the non-breeding season, he or she will continue on the same annual cycle for the rest of his/her life. One of the big unknowns for waders/shorebirds – and for other groups of migrant birds for that matter – is just how these settlement patterns develop. (See Generational Change to read how young birds can create new patterns of migration).

Checking their data, Jeroen and his colleagues could see no pattern in the juvenile/adult proportions, sex ratios or sizes of the birds in different non-breeding areas that would help to explain differences in fitness. Birds from the same breeding areas of Greenland end up in non-breeding locations along the whole north-south range. It is almost as if the dice are rolled and a juvenile ends up where chance events take it.

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At the level of the individual:

  • A Sanderling that spends the non-breeding season in Ghana does not know that it would have a better annual survival rate and be likely to return earlier to Greenland each spring had it ended up wintering in England or travelled as far as Namibia.
  • A bird in Namibia has no idea that it could have saved itself an accumulated migration distance of 37,000 km each year by stopping in England, without affecting its probability of survival.
  • A first-year bird that spends its first potential breeding season feeding on the beaches of Mauritania, will be unaware that first-years from Portugal have all travelled north to Greenland.

blog 8 look for crIt’s amazing what colour-ring readers have helped to discover but there is much still to learn about the migration strategies of individual waders.

Let’s hope that birdwatchers will continue to look out for colour-rings, as flocks of Sanderling chase the waves in and out on beaches throughout the world.

 

Paper in Journal of Animal Ecology

Low fitness at low latitudes: wintering in the tropics increases migratory delays and mortality rates in an arctic- breeding shorebird

Jeroen Reneerkens, Tom S. L. Versluijs, Theunis Piersma, José A. Alves, Mark Boorman, Colin Corse, Olivier Gilg, Gunnar Thor Hallgrimsson, Johannes Lang, Bob Loos, Yaa  Ntiamoa-Baidu, Alfred A. Nuoh, Peter M. Potts, Job ten Horn & Tamar Lok.

blog 6 run for it


GFA in Iceland

Graham (@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.

Sixty years of Wash waders

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Weighing a Turnstone

The Wash Wader Ringing Group (WWRG) started with a bang on 18 August 1959, when the team made a catch of 1,132 birds in a Wildfowl Trust rocket-net at Terrington, in Norfolk. Over the years, cannon have replaced rockets, catches have become generally smaller and the scientific priorities have been refined, but the Group continues to focus upon discovering more about the waders that use the Wash. This blog attempts to summarises what has been learnt about the waders that rely upon the Wash, the vast muddy estuary that lies between Lincolnshire and Norfolk, on the east coast of England.

Sixty years ago, the first goal was to understand where the vast flocks of waders that visit the Wash came from – a task that would provide great insights into the way that the whole East Atlantic Flyway works. In this time, over 300,000 birds have been caught and ringed on the Wash, as you can see in the table below. Equally importantly, hundreds of bird-ringers from across the UK and scores of visitors from around the world have joined WWRG teams, in order to learn more about the study of shorebirds. Further international collaboration has been fostered through overseas visits by WWRG members and emigration of some key personnel. The impact of the Group is truly global, as you can read in the WWRG report for 2014/2015.

wwrg table

A total of 307,226 birds is impressive, especially when some of the species totals are compared to the national totals of the BTO Ringing Scheme for the whole of Britain & Ireland since 1909. WWRG is responsible for over 40% of the Grey Plover, Knot, Sanderling and Bar-tailed Godwit, with Grey Plover topping the list at nearly 60%. These are terrific achievements for a group of volunteers. I don’t have the figures but I reckon that Nigel Clark has been responsible for the largest number of catches.

Wee quiz: What’s the best match between these Wash waders and the countries that they are quite likely to have come from? Answers at the end of the blog:

  • Species: Bar-tailed Godwit, Black-tailed Godwit, Curlew, Oystercatcher, Sanderling, Turnstone
  • Countries: Canada, Greenland, Iceland, Norway, Finland & Russia
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Firing box connected to 4 cannon-nets

In the early days, rocket nets were borrowed from the Wildfowl Trust for an annual summer week of catches, but the development of cannon-nets gave opportunities for all-year ringing. The intensity of the Group’s activities grew in the 1970s, when there was a threat to build a freshwater reservoir on the mudflats. For a couple of years, Clive Minton (founder and leader) persuaded us to visit fortnightly, so that we could get better data on weight-gain and turn-over, using a mixture of cannon-netting and mist-netting. Everything we knew was published by the Group as The Wash Feasibility Study in 1975. These days, the Group gets together about ten times a year for catching and colour-ring-reading sessions.

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By catching and ringing large numbers of the key species that visit the Wash, the Group was able to generate maps showing what are now well-known patterns of migration (see Which wader, when and why?). Early on in the Group’s history, there was a focus on nine species, with Black-tailed Godwit added as a tenth when numbers increased. Each of these species has its own section below. The maps were prepared for the Wash Wader Ringing Group 2016/2017 Report by Ryan Burrell, using data stored within the BTO archives. Blue dots represent WWRG-ringed birds that have been found abroad. Red triangles represent foreign-ringed birds caught on the Wash. The base maps used are by courtesy of Natural Earth (www.naturalearthdata.com).

Oystercatcher

wwrg map OCThe map alongside clearly demonstrates the strong link between the Wash and Norway. Other interesting things that have been discovered about Oystercatchers:

  • They live a long time. An Oystercatcher that we caught at Friskney on 30 July 1976 broke the longevity record for a BTO-ringed wader when it was shot in France on 4 April 2017 (41 years 1 month and 5 days). It was ringed as an adult so we don’t know the exact age – but it must have been at least 43 years old. There’s a WaderTales blog with a list of longevity records for BTO-ringed waders.
  • When life gets tough, Oystercatchers fail to complete their autumn moult, retaining some of their outer primaries for an extra year. The ability to complete moult and annual survival rates are both affected by cockle and mussel supplies on the Wash. There’s more about this in two papers in Biological Conservation and the Journal of Applied Ecology.

Grey Plover

wwrg GV GVIn the early days of the WWRG, Grey Plovers occurred in much smaller numbers than they do now. Writing in an article about the first 40 years of the Group, Clive Minton told the story of the first catch of 100, made in 1963, that was celebrated with three bottles of champagne provided by the late Hugh Boyd, delivering on an incentive that he had promised.

  • Over half of the Grey Plover that have been ringed in Britain & Ireland since 1909 have been ringed by WWRG since 1959 (58.9%)
  • All of the Grey Plover using the Wash breed in Siberia. Some birds spend the winter on the Wash but there are autumn moulting flocks of birds that will go on to winter in other parts of Britain & Ireland, and spring and autumn passage of birds that travel as far south as West Africa.
  • Grey Plover are late to leave the Wash, with the last departures not occurring until the start of June. Unsurprisingly, they are some of the last waders to return at the end of summer, which puts pressure on birds to finish moult before the short, cold days of winter. Some adults fail to complete primary moult, especially if food supplies are low. There is more about Grey Plover moult in this WaderTales blog.

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Knot

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First-winter Knot (subterminal bands on wing coverts and, as yet, unmoulted juvenile fethers on upper-parts)

Knot (or Red Knot) are truly international waders, as is shown in this map of movements of islandica  (and a few canutus) birds  to and from the Wash. Several WWRG members have been heavily involved in efforts to understand the decline in numbers of the rufa subspecies in Delaware Bay (on the North American eastern seaboard) and Clive Minton has been at the heart of efforts to explain the sudden drop in survival of piersmai and rogersi adults that winter in Australia and migrate to Arctic Russia via the Yellow Sea (see Wader declines in the shrinking Yellow Sea).

  • We are still learning about Knot migration. The cluster of reports of WWRG-ringed birds in Northern Norway looks odd on this map projection but it turns out that this is a well-used stopping-off point for islandica Knot heading for northern Greenland and NE Canada. This route was first confirmed in 1985, when a joint Durham University and Tromsø University expedition caught 18 Wash-ringed birds in a total catch of 1703 birds.
  • wwrg net set

    Setting cannon-nets

    Many birdwatchers visit the Wash in autumn and winter to see the swirling Knot flocks at Snettisham and Holme. If high tide is at first light, Knot and other waders sometimes roost on Heacham Beach, giving the occasional opportunity to make a significant catch. The most recent of these, on 11 February in 2012, included 2757 Knot, 77 of which were already wearing rings.

  • The most recent analysis of wader populations in Great Britain showed that there was a drop of nearly 20% in wintering Knot numbers (from 320k to 260k) in less than a decade (see Do population estimates matter?). Regular catches on the Wash will help produce estimates of annual survival rates and age ratios of the islandica subspecies.

Sanderling

wwrg sanderlingThe biggest catches of Sanderling are generally in the summer, when the Wash is a meeting point for birds from Greenland and Siberia. July can sometimes see catches of 200 or more birds. Traditionally, a Sanderling catch was the curtain-raiser at the start of Wash Week, an opportunity for the whole team to make one catch before splitting into ‘Terrington’ and ‘Lincolnshire’ teams for the rest of the main summer trip.

  • Wintering Sanderling on the Wash are thought to be exclusively of the race that heads northwest in the spring, to Greenland via Iceland.
  • Late summer and spring see the addition of birds passing through on their way from/to Siberian and Greenlandic breeding areas.
  • I well remember the first time we caught a Sanderling (on 26 July 1975) wearing an Italian ring (caught in Italy 9 May 1975). Thanks to Jeroen Reneerkens (whose work will be covered in an upcoming blog) I now understand that this is probably a bird that migrates from Namibia to Greenland in spring, via the Mediterranean. It will have been on its way back to Namibia when caught in July.

wwrg map SS DN

Dunlin

wash dunlin

Sam Franks, looking for the buffy tips on inner coverts, which distinguish first-year birds from adults

Nearly half of the waders caught by WWRG have been Dunlin – a total of 140,168 up until the end of 2018. There were really big flocks of Dunlin in the 1970s but numbers have dropped over the years, with peak counts now half what they were, according to WeBS data.

  • We caught over 3,500 Dunlin in one week in 1976 but the annual total has exceeded 1,000 in only four of the last ten years. Partly, this reflects a change in behaviour in the summertime, with fewer waders roosting on fields and hence less catchable.
  • Three races of Dunlin visit the UK. Our winter birds are alpina, from Siberia, NW Russia and northern Scandinavia. A lot of July birds are schinzii, breeding in the UK and as far north as Greenland, and we occasionally try to convince ourselves that we have caught an arctica from northern Greenland.
  • Data collected for the WeBS survey suggest that national winter totals have dropped by over 40% in 25 years. This could perhaps partly be explained by a redistribution of alpina, with new generations of young birds settling in wintering areas on the other side of the North Sea. Warmer winters may well make this a more practical proposition than in the 1970s. There’s more about this in this paper.

Black-tailed Godwit

wash blackwit

Newly ringed Black-tailed Godwit, caught in a mist-net at night.

Black-tailed Godwits became a priority species in 1995, when Jennifer Gill (University of East Anglia) started a project to study the movements of individuals, using colour-rings. Nearly 25 years later, the WWRG-ringed Black-tailed Godwits have contributed data to numerous papers, largely focusing upon migration.

  • The Wash is a hugely important area for moulting islandica Black-tailed Godwits. Some birds stay in East Anglia for the winter but others move south and west within the UK, west to Ireland and south to France, Portugal and Spain.
  • There are several blogs about Black-tailed Godwits in this WaderTales contents list.

Bar-tailed Godwit

One of the key things that was learned from the sudden decline in annual survival rates in a range of species that use the Yellow Sea (as mentioned above) is a need for regular monitoring of marked birds. The WWRG’s Scientific Committee set up colour-flagging projects for Bar-tailed Godwit, Curlew and Grey Plover, in order to increase the reliability of estimates of annual survival for three species that the Group does not catch in sufficient numbers to generate good retrap histories. Birdwatchers can help by reporting colour-marked birds here.

wwrg barwit map etc

  • In Bar-tailed Godwits: Migration & Survival there is a comparison of the data generated by a catch of 505 Bar-tailed Godwits in 1976 with the information that has been generated recently, using colour-flags.
  • Bar-tailed Godwits are long-lived birds. A WWRG bird holds the current record for a BTO-ringed Bar-tailed Godwit: 33 years and 11 months between ringing in 1978 and recapture in 2008. BTO longevity records are discussed in this WaderTales blog.
  • Colour-ring reading is now a significant element of Group activities, as described by Rob Pell in the WWRG Report for 2016/2017.

Curlew

Back in the 1970s, Curlew were still hunted on the Wash (paté made from autumn-shot birds was reported to be very tasty). Shooting stopped in Great Britain in 1981, when the maximum winter count on the Wash had dropped to about 3,000 birds, and by 2003/04 the maximum winter count was 15,336. Since then, numbers have declined, in line with national and international trends.

wwrg curlew map etc

  • A large number of Curlew on the Wash in winter are from Finland and surrounding countries. Surprisingly few are of UK origin.
  • Birds wearing WWRG leg-flags have been observed breeding in the Brecks (Norfolk/Suffolk).
  • The Curlew is internationally designated as ‘Near Threatened’. Is this really true when we can still see a field with 1000 roosting Curlew in Norfolk? Answers here.

Redshank

wash redshThe latest population estimates suggest that Great Britain has lost 26,000 wintering Redshank in less than a decade, representing a drop of 20%. Perhaps WWRG data can be used to help to explain these declines? Here are some of the things we know:

  • The Redshank on The Wash in the winter are mainly a mixture of birds from around the Wash, across the UK and from Iceland.
  • In cold winters, Redshank wintering on the Wash die in large numbers. After a period of severe weather in 1991, nearly 3,000 wader corpses were collected from along the tide-line, about 50% of which were Redshank. The winter WeBS counts for Redshank dropped by 50% after this mortality event but have recovered somewhat since then.
  • An analysis of nearly 1,000 dead Redshank showed that about two-thirds were of Icelandic origin. There was a tendency for smaller birds to be more susceptible to cold weather mortality than larger birds of the same species (More information in this paper by Jacquie Clark)

wwrg map RK TT

Turnstone

wash ttWinter Turnstone are birds that will head for Greenland and NE Canada in the spring but recoveries of birds in Finland and other Scandinavian countries indicate a passage of continental birds. African recoveries of WWRG-ringed birds probably include birds from Canada/Greenland and Finland/Scandinavia.

  • Turnstone wearing US Fish & Wildlife Service rings are occasionally caught on the Wash. Some of these rings were put on by Guy Morrison and his colleagues in Alert, Ellesmere Island, Canada. Guy was an early member of WWRG. It’s a small world!
  • The first Wash Turnstone were colour-ringed in 1999, as part of a study to understand why birds were feeding on the docks at Sutton Bridge. There is a WaderTales blog about the resulting paper by Jen Smart and Jennifer Gill. Colour-ringing continues, to measure annual survival rates.
  • Turnstone have a reputation for eating almost anything (including dog excrement and a human corpse) so do not be surprised if you see a colour-ringed bird scavenging for chips on the Hunstanton sea-front.

A few more highlights

Ringed Plover: this is not one of the ten key study species but 1,432 have been ringed between 1959 and 2018. Some birds are local breeders that hardly move anywhere but other birds link the Wash with Greenland, northern Norway, Morocco and Senegal.

wwrg GKGreenshank: The Group supports a colour-ringing project that was initiated by Pete Potts, in Hampshire. More information here.

Spotted Redshank: During the period 1959 to 2018, WWRG ringed a total of 85 Spotted Redshank, representing over 20% of the total ringed in Britain and Ireland since 1909. Amazingly, sixty of these birds were ringed on the same day – 27 July 1975. There is a blog about this catch and the recent decline in the number of Spotted Redshank visiting the UK. Fewer Spotted Redshanks.

Ruff: Until its closure, WWRG members spent many a smelly night at Wisbech Sewage Farm. This was a great place to catch Ruff, Curlew Sandpipers, Green Sandpipers etc. in mist-nets. Group members wrote a paper about Ruff moult and migration.

Rares: Occasionally there are surprises! WWRG has caught one each of Stone Curlew, Pectoral Sandpiper, Broad-billed Sandpiper and Terek Sandpiper. The last bird features in this WWRG blog.

What do we know now?

Migration studies have revealed the importance of the Wash to half a million or more waders each year – birds that spend the whole winter, others that refuel in the spring and vast numbers that rely on the food supplies in the mud to provide the energy for the post-breeding moult. There’s a selection of papers that have included WWRG data here, on the Group’s web-site.

wwrg cr TTThe Group still aims to maintain its general ringing programme, so that a representative sample of the key species carry rings. Colour-ringing projects aim to provide survival estimates for Curlew, Bar-tailed Godwit, Grey Plover and Turnstone, with Greenshank and Black-tailed Godwit colour-rings contributing to migration studies. Birdwatchers who visit the Wash can help by reporting colour-marked birds here, on the WWRG web-site.

WWRG data have been used to help inform decisions about the future of the Wash but the threats keep coming. Studies of migration and seasonal turn-over in numbers contributed hugely to decisions to provide national and international protection to the area and to fend off the 1970s plan to build a freshwater reservoir on the rich mudflats. The information that has been generated by many generations of volunteers over a period of sixty years has been used to manage the level of shellfish exploitation, to inform decisions about wind turbine locations and to manage activities that can cause disturbance.

The Wash Wader Ringing Group is very keen for its data to be used – and not just for impact assessment studies. Click here to learn more.

Diamond Jubilee

PLI

Phil Ireland releasing a Curlew

Over one thousand people are estimated to have contributed to sixty years of the Wash Wader Ringing Group’s activities. We have lived in barns, rolled cars, dug tens of thousands of holes, carried nets for miles, made important catches, had depressing failures, got frostbite, been threatened by surge tides and made friends for life.

In the whole of this period, there have been only two leaders of the Group – Clive Minton (1959-1981) and Phil Ireland (1981-present). Bird ringers, wader biologists and millions of waders owe them both a huge debt of gratitude.

You can read more about the history of WWRG on the Group’s website:

wwrg sunset

Photo at the top of this blog is by Cathy Ryden. Many thanks to her and to other photographers.

Wee quiz:

  • Bar-tailed Godwit – Russia
  • Black-tailed Godwit – Iceland
  • Curlew – Finland
  • Oystercatcher – Norway
  • Sanderling – Greenland
  • Turnstone – Canada

 


GFA in Iceland

Graham (@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.

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Generational change

blog TGG on postIn a changing world, with more chaotic weather patterns and rapidly altering habitats, migratory birds are faced with opportunities and challenges. Long-term monitoring of colour-ringed Black-tailed Godwits, during a period of range expansion and phenological change, has revealed that individuals behave consistently over time but that the behaviour of new generations is moulded by the conditions they encounter.

A changing world 

When trying to explain observed changes in the distributions and annual cycles of migratory birds, there are many things to consider:

  • blog VM y flag

    Colour-rings enable life-time tracking. This bird, caught on its nest, had been ringed as a chick.

    Are individual birds able to take advantage of new breeding and non-breeding sites, as they become available, particularly if other areas become less suitable?

  • Are individuals able to change the timings and patterns of migration?
  • Do individuals adjust their migration routes as a consequence of changes in stop-over or wintering areas?
  • If individuals do not change what they do, how do we explain range expansions and changes in timing of migration?

Put simply, how does climate change lead to changes in distribution of migratory birds? Answering this question is key to being able to predict the rate and direction of future changes, and to assess whether our existing networks of protected sites will continue to support populations in the way that was intended. This issue was tackled by Jennifer Gill, José Alves and Tómas Gunnarsson in their paper “Mechanisms driving phenological and range change in migratory species”, published in Linking behaviour to dynamics of population and communities: applications of novel approaches in behavioural ecology and conservation, a special issue of Philosophical Transactions B (Royal Society).

Potential models

Change could happen in two main ways:

  • Individuals could relocate – having knowledge of a range of available conditions, they can choose to move elsewhere.
  • New generations could settle in new areas (in the breeding season, the non-breeding season or both) and/or adopt new migratory strategies.
blog map

Map that illustrates range expansion

Working out whether change happens through individual movement or generational shifts can only be done by life-long tracking of individuals, in populations in which range change is happening. The Icelandic population of Black-tailed Godwit is ideal for such an investigation. Black-tailed Godwits have been expanding into new breeding areas of Iceland for over 100 years, as discussed in this WaderTales blog. Population growth has been facilitated through warming spring conditions, as discussed in From local warming to range expansion.

blog TGG juvs

Naive youngsters, gathering together before migration

Icelandic Black-tailed Godwits fly south in the autumn, to spend the winter in the British Isles, The Netherlands, Belgium, France, Spain and Portugal. As numbers have grown, winter counts have increased in many areas, with new flocks appearing and expanding on estuaries and areas of wet grassland where the species was previously absent or scarce.

 

Winter distribution

The Wetland Bird Survey shows that there are three times as many Black-tailed Godwits wintering in Great Britain as there were 25 years ago. The biggest changes in numbers have occurred on estuaries in the northwest of England, with the Morecambe Bay winter maximum rising from about 180 to 3200, for instance. Where have these extra birds come from?

blog juvs on Axe

These young birds happen to have ended up on the Axe Estuary in Somerset

Black-tailed Godwits have been ringed in Iceland for nearly twenty years, providing a pool of known-age adults for which natal sites are known. Winter observations of colour-ringed individuals have shown an interesting pattern; birds breeding in newly-colonised areas, particularly in north and east Iceland, are the ones that are more likely to be found in newer winter sites.

In their paper, the authors suggest that birds nesting in these colder areas, where spring comes later, will be fledging quite late and leaving Iceland after adults have departed. With no experienced birds to follow, these young birds may well stop off at the first suitable site, many of which are in the north of the wintering range, and then they return to breed in their natal sites. Birds in Morecambe Bay don’t know that days are longer and the weather is kinder for other birds that travel further south to wintering areas such as Portugal.

blog RS Dee

Wintering birds in Northwest England

Observations from birdwatchers show that the same colour-ringed individuals are nearly always found at the same wintering sites each year. Whatever mechanism is producing this new-breeding-site to new-wintering-site link, it is becoming clear that older birds continue to do what they have always done, with changes in distribution happening as a result of a generational shift.

The annual cycle

Colour-ringed Black-tailed Godwits have been tracked for over 25 years, with a small number of individuals contributing data for the whole of this period. This tracking information can be used to ask how much individuals move around and experience different sites and to assess whether individuals from different generations are using different parts of the range.

Using colour-rings, the Black-tailed Godwit team has discovered that, although individuals can live for over 20 years, in that time they generally use a total of only about four sites between leaving Iceland in late summer and returning in the spring. Basically, individual birds have very limited experience of sites and there is no evidence that they have moved to occupy different sites as, for instance, winter conditions have changed.

blog infographic

Spring arrivals in Iceland

Colour-ring observations have shown that individual birds do not change their breeding or wintering locations and that migrating individuals often appear in the same stop-over sites year after year. The timing of movements is also pretty consistent, especially in the spring. A previous WaderTales blog called Why is spring migration getting earlier? demonstrated that the timing of  migration of individual Black-tailed Godwits varies very little, with observed shifts in the period of migration being driven by young birds returning to Iceland for the first time on average doing so somewhat earlier than previous generations. Once individual birds settle into a timing pattern, they stick to it.

blog LJ arrivals

Black-tailed Godwits, newly arrived in Iceland after crossing the Atlantic

Migration patterns

As discussed above, individual Icelandic Black-tailed Godwits have experience of only a small number of sites, which they use on an annual cycle. When migrating, a bird will generally use the same stop-over site when breaking its journey south, to undertake autumn moult, or on their way north, to take on fat for the trans-Atlantic journey. There is a range of spring migratory strategies in islandica Black-tailed Godwits, as discussed in Overtaking on migration.

blog wwrwOnce established, the annual migratory programmes of individuals rarely change, as illustrated by the map to the right. Colour-ringed Black-tailed Godwit W-WR/W regularly moulted on the Wash, in eastern England, before spending the late winter and spring in northwest England. In the late summer of 2002 he was reported at Slimbridge on 18th and 20th July but back on the Wash on the 25th. Having made the Atlantic crossing and ended up in southwest England, he was able to correct what he may have perceived to be his mistake, returning to the moulting area that he had been using since at least 1996.

Individuals might not change their annual migration routes but we do see changes in numbers on different sites that are used during migratory stop-overs. In a paper published in 2018, Mo Verhoeven and colleagues investigated whether observed changes in migratory patterns of a population of the limosa subspecies of Black-tailed Godwit were caused by individuals altering their strategies or by generational change.

Limosa Black-tailed Godwits leave breeding areas in countries such as The Netherlands in late summer, heading south to either West Africa or Iberia, where they spend the winter. In spring they all gather in staging sites in Portugal and Spain, typically on rice fields. Over the course of less than ten years, the average peak number in Extremadura (Spain) has dropped from about 24,000 to 10,000, while the numbers on the Tagus and Sado estuaries rose from 44,000 to 51,000. These changes took place during a period of rapid population decline, as described in this blog focusing on a paper by Rosemarie Kentie and colleagues.

blog VM Tagus

Limosa Black-tailed Godwits feeding in a rice field in the Tagus estuary

Mo Verhoeven et al have shown that this rapid population-level shift in spring stop-over sites from Spain to Portugal, 300 km further west, was driven by young godwits increasingly using Portugal in the period January to March, instead of Spain. Nearly all of the older birds stuck with the routes they knew. The paper is Generational shift in spring staging site use by a long-distance migratory bird.

Change happens to birds

One thing that is becoming clear in Black-tailed Godwits is that birds are being affected by change – individuals do not have the knowledge or flexibility to effect change. Even in long-lived birds, like Black-tailed Godwits, we see no evidence of individuals altering what they do over what is now two decades, despite the fact that the species’ migration dates, wintering areas and migration routes have all perceptibly changed over the same time period. It’s all about generational change. The behaviour patterns of young birds arise from the conditions they encounter in the first year of life, after which they are repeated.

Details of the Generational Change paper by Gill et al

blog LJ sum plumThe paper at the heart of this blog is: Mechanisms driving phenological and range change in migratory species by Jennifer Gill, José Alves and Tómas Gunnarsson, from the Universities of East Anglia (UK), Aveiro (Portugal) and Iceland. It is published in Linking behaviour to dynamics of population and communities: applications of novel approaches in behavioural ecology and conservation, a special issue of Philosophical Transactions B (Royal Society).

The paper could not have been produced without the help of “thousands of observers of colour-ringed godwits who have made these analyses possible”. This WaderTales blog is a celebration of the work they do: Godwits and Godwiteers.


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

 

Time to nest again?

blog Snipe TGGEarly return to breeding areas is widely acknowledged to be ‘a good thing’ but why? Some people suggest that early migrants can choose the best territories, others argue that early chicks have a disproportionately high chance of fledging but there are other explanations too. In their paper in Ecology & Evolution, Catriona Morrison and her colleagues ask how much of the advantage of being an early migrant could be associated with having an option to nest again, if the first attempt fails.

Setting the scene

In a previous WaderTales blog, about Icelandic Black-tailed Godwits, there is clear evidence that the species is increasing in number and spreading into new breeding areas. In another blog you can read that the expansion is linked to warmer spring conditions, which allow earlier nesting. How might this change in nesting phenology influence overall productivity and contribute to the population growth in Black-tailed Godwits, and do the same processes work for other species?

blog oyc godwit

Individual Black-tailed Godwits that arrive in Iceland early each year may have a higher chance of nesting successfully, just because they have time to try again if the first nest fails

In their 2019 paper, Catriona Morrison and her colleagues from the Universities of East Anglia (UK), Iceland and Aveiro (Portugal) used a simulation model to ask whether the greater time available for laying replacement clutches can create a pattern of increased productivity among early-arriving migrants, without the need to think about territory choice or local resource availability. They suggest that early arrival can lead to greater breeding success simply because early birds have more time available to nest again, following nest loss. Within the model they explore the effect on breeding success of varying several important relationships:

  • blog Snipe nest

    This early Snipe nest might get predated but there should be time to try again

    Whether early clutches are more likely to hatch than later ones (seasonal variation in nest survival rates) – such a trend could be created by predation patterns, resource availability and opportunities to conceal nests.

  • Whether one or more replacement clutches is possible within the time available during the breeding season (number of re-nesting attempts).
  • Whether late chicks are less likely to survive and become breeding adults than earlier ones (seasonal variation in recruitment rates) – which would make re-nesting attempts less valuable.

Results

The models developed for the paper showed that, when the chance of losing a breeding attempt does not change during the course of the breeding season, species experiencing intermediate nest survival rates will benefit most from re-nesting. This makes sense; a species that has a very high chance of hatching its chicks will not need to re-nest and one that has a negligible success rate is not going to do much better if it lays more than one clutch.

blog Lapwing

This late-nesting Lapwing may not be able to defend its nest

Nest success may not be constant over the course of a season. Late pairs may find it harder to distract predators if they don’t have the support of other breeding birds, with a consequent drop in success over the summer. Alternatively, species that nests in clumps of grass, such as Snipe, might find it easier to hide their nests later in the season, thereby increasing nesting success over time.

Picking out just a few of the scenarios that are covered in more detail in the paper:

  • When nest survival rates are constant and replacement clutches are possible, early arrival increases the probability of achieving a successful nesting attempt. These benefits of early arrival can be substantial enough to persist even when late-hatched chicks (from replacement nests) are less likely to survive and recruit into adulthood.
  • If there is a seasonal decline in nest survival, late-arriving individuals will have far fewer successful nesting attempts in their lifetime than early-arrivers. In this case, laying replacement clutches only slightly increases the number of successful nesting attempts and the subsequent number of recruits.
  • If there is a seasonal increase in nest survival, early-arriving individuals will tend to lose their first clutches but these individuals have time to re-nest, and are likely to fledge the subsequent attempt. Late-arriving individuals arriving will be more likely to have a successful first attempt and hence the number of successful nesting attempts varies little with arrival date.

The main take-home message of the paper is that, in almost all of the circumstances considered, early arrival can lead to higher breeding success, simply because of the greater time available to lay replacement clutches.

Blog RP migration

What does this mean for waders?

blog Oyc nest

An Oystercatcher does not need much time to ‘build’ a nest

Repeat nesting is a common strategy in waders; a female Oystercatcher, for instance, can quite quickly lay a second clutch if the first clutch is lost. Strategies exist that can lead to a female having more than one successful brood in a season, as seen when a female Dotterel leaves a male to incubate a clutch of eggs and moves on to another male. In most circumstances, however, a pair of waders has time to raise one brood of chicks in a season, by succeeding with the first attempt or taking opportunities to lay replacement clutches if time and resources allow.

It is obvious that, if nesting success is very high, there will be little need to lay a second clutch and if success is really low, little will be achieved by laying more clutches. Waders tend to have intermediate nest-success; most are ground-nesters, making them vulnerable to a wide range of mammalian and avian predators of eggs and chicks. The scenarios modelled in the paper are particularly (but not exclusively) appropriate to breeding waders

blog Knot

Knot – a High Arctic breeder, constrained by a short season

The modelling used in this paper shows that having the time to try again is likely to increase the probability of annual success, as long as the breeding season is long enough. We know that pairs of Ringed Plovers breeding at temperate latitudes have time for several breeding attempts but pairs at high latitudes may have little chance for a second attempt, especially if nest failure occurs late in the incubation period. One way of increasing the time available to breed is to arrive earlier and the benefits of early arrival may be particularly strong for birds that occupy areas where there is a lengthening potential breeding season, something that can be made possible through climate change and warmer springs.

It is not uncommon for a breeding wader to live for five years, ten years – or even longer for larger species (WaderTales blog). During its lifetime, an individual may experience breeding seasons with differing levels of predator activity or other causes of nest loss, such as flooding or trampling, might occur. Although an individual might migrate at the same time each spring, the number of nesting attempts it will be able to fit in during any particular year will depend upon factors such as weather, prey availability and predation pressure.

blog Sanderling

Sanderling with chicks

Nest survival rates in wader populations can show seasonal declines (e.g. Sandercock 1999 – Semipalmated Sandpipers), increases (e.g. Reneerekens et al. 2016 – Sanderling) or little seasonal variation in survival (e.g. Sandercock 1999 – Western Sandpiper), but in all cases there is variability between years. All of these seasonal patterns of survival change were modelled in the Morrison et al paper. In almost every situation, a wader will have a higher chance of successfully rearing youngsters if it (and its mate) are on an early spring migration schedule.

Summary

blog Oyc

This Oystercatcher may regret nesting early! If it fails, it can try again.

Turning up early on breeding grounds in spring can potentially lead to higher reproductive success, solely as a result of the greater time available for laying a replacement clutch. Using modelling, Catriona Morrison and colleagues show that this early-arrival-benefit can be conferred even when later nesting attempts are less likely to produce successful recruits.

Advances in the timing of spring migration are occurring in many species and these findings highlight the potential role of replacement nests as a driver of population increase in those areas where repeat nesting becomes increasingly possible. Professional ornithologists and citizen scientists who study nesting birds (not just waders) are encouraged to do so for the whole season, especially by following marked individuals. Birds that wear geolocators, which can record incubation patterns for nesting attempts that would otherwise remain undetected, may be particularly helpful when trying to discover just how likely birds are to re-nest and with what success.

Only part of the story

Blog tag

Geolocator on Whimbrel

In the long run, the success of an individual bird can be measured by the number of offspring it has in its lifetime and even by the number of its genes that are present in future generations. The number of chicks that fledge each year is only part of the story, therefore. How many of these youngsters recruit to the breeding population? Do they end up breeding in areas where they will have high breeding success? Will their progeny live for a long time and hence have many opportunities to produce their own chicks? Long-term wader studies might reveal some of these answers – eventually.

Paper

blog RP chicks

The aim: a successful brood

The paper was published in Ecology & Evolution.

Why do earlier-arriving migratory birds have better breeding success? Catriona A. Morrison, José A. Alves, Tómas G. Gunnarsson, Böðvar Þórisson and Jennifer A. Gill.

DOI: https://doi.org/10.1002/ece3.5441

The paper is freely available to view.

 


GFA in Iceland

Graham (@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.

Whimbrel: time to leave

blog WW-WLGeolocators* have provided fantastic information about the movements of migratory birds – making links between countries, revealing previously unknown stop-over sites and indicating just how quickly birds traverse our planet. A small number of Icelandic Whimbrel have carried geolocators for up to six annual cycles, providing Camilo Carneiro with an opportunity to investigate the annual consistency of egg-laying, autumn departure, arrival in West Africa, departure in the spring, stopover in Western Europe and arrival back in Iceland.

* Geolocators are tiny devices that record the daily positions of birds, by measuring the timing of dawn and dusk. An individual typically carries a geolocator for a year and then needs to be re-caught for the data to be downloaded.

Planning a trip

When booking a train journey on-line, the first question I am asked is whether I want to stipulate departure time or arrival time.  In early spring, with breeding on their minds, you might think that Whimbrel will focus on the time they need to be in Iceland, rather than the time they leave West Africa? If that’s the case then it might be best to take early spring opportunities if they arise, to catch express winds that will make the journey as rapid as possible and to get to Iceland early. Is that the case?

blog mangroves and beach

The Whimbrel is one of several wader species that breed in Iceland. Each autumn, Redshank, Snipe, Golden Plover, Oystercatcher and Black-tailed Godwit fly south to Europe, especially Ireland and the United Kingdom, but many Ringed Plover, most Dunlin and most Whimbrel travel as far as Africa. The main wintering sites for Whimbrel are in West Africa, south of the Sahara, in countries such as Guinea-Bissau. Here they can be seen feeding on crabs on the mangrove-fringed muddy shoreline (above). It’s a very different environment to the inland floodplains of Iceland (below).

blog river plain

In a paper by Tómas Gunnarsson & Gunnar Tómasson in 2011, we learnt that Whimbrel arrival times in Iceland did not change much between 1988 and 2009 (just 0.16 days earlier per year), while timing of arrival was advancing much more in species that travel less far to winter grounds, as you can see in this diagram.

wader arrival Tand G

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

The arrival date for Black-tailed Godwit was advancing fastest (0.81 days per year). In more recent research, it has been shown that the rapidly advancing trend for Black-tailed Godwits is being driven by new recruits to the population – individual adults are not changing their schedules. Why is spring migration getting earlier? summarises a paper by Gill et al in Proceedings of the Royal Society B.

The Whimbrel trend has been recalculated, with a longer run of years, and the new change of 0.03 days earlier per year is not significantly different from zero. Given that Whimbrel are breeding alongside other species that are arriving in Iceland much earlier than thirty years ago, what are the constraints to the timing of their migrations?

Migration timings for Whimbrel

Camilo Carneiro, José Alves and Tómas Gunnarsson from the Universities of Aveiro (Portugal) and Iceland have been studying a population of Whimbrel in Southern Iceland. Birds are caught on the nest in one year and then re-caught in the subsequent year – or two years later if a bird evades capture in the intervening summer. The following results summarise weeks and weeks of patient fieldwork and brush over the hours of frustration caused by wary birds that have been caught before!

blog catching

Over the course of the whole study, 86 Whimbrel were fitted with geolocators, 62 of which were retrieved. Repeatability could be calculated for 16 birds, with between 2 and 7 years of data collected from each individual. The results are summarised in these few bullet points. Please see the paper for confidence intervals and more details about differences between the sexes.

  • Blog tagIndividual timings of autumn departure from Iceland varied between years. The repeatability index is 0.28, with a suggestion of a gender difference (females 0.40, males 0.02). Males tend to look after chicks for a longer period than females so their departure dates may be more strongly influenced by the success of each year’s breeding attempt.
  • Autumn arrival time in West Africa was closely linked to departure time because, on all but one occasion, southward migration was achieved through a single direct flight. See Iceland to Africa non-stop.
  • Spring departure time from West Africa was highly consistent, with a repeatability index of 0.76 and no discernible difference in repeatability between males and females.
  • blog long green grassSpring arrivals in Iceland. Some Whimbrel that managed to complete spring migration in a single flight in some years stopped off in other years. These breaks, perhaps to wait for more helpful wind conditions and/or to refuel, resulted in variability in annual arrival dates for individuals. The repeatability for the two sexes combined was 0.23.
  • Laying date was the least consistent stage of the annual cycle, with a repeatability index of 0.11 and no significant difference between males and females.

In an individual Whimbrel’s annual cycle, there appears to be one fixed point – departure from wintering ground in West Africa. With no discernible seasonality of resource availability on the wintering grounds and little change in day length in these areas, departure dates are probably being determined by an ‘internal clock’. Two major unknowns will then determine what happens in the next twelve months. Will wind and weather conditions be conducive to a one-leg flight to Iceland and how successful will a bird be in any particular breeding season? Unforeseen events, such as having to wait for a delayed partner, losing a first clutch, and the time needed to guard chicks will all affect the timing of autumn migration.

Understanding individuals

blog tag through grassThe study of wader migration has advanced hugely.

  • Fifty years ago, the main measure of migration phenology was the appearance of the first individuals of a species.
  • Colour-ring sightings are ideal for providing repeat arrival dates over the lifetimes of individuals, as exemplified by the Gill et al paper on Black-tailed Godwits, which suggest that individual timing is highly repeatable.
  • Geolocators have provided more detailed information about the precise arrival and departure timings of individuals, which is so important if we wish to conserve threatened, migratory species that visit areas in which data collection was previously virtually impossible.
  • Now, by tracking individual birds for several years, it is possible to look at the variability in annual patterns, and what can cause this variability.

Over the next decade or so, as devices get smaller and remote downloads become easier (eg using GSM tags), it should become possible to understand the conditions that lead to fast, slow and aborted migratory journeys in a whole range of species. Exciting times!

Paper

Why are Whimbrels not advancing their arrival dates into Iceland? Exploring seasonal and sex-specific variation in consistency of individual timing during the annual cycle. Camilo Carneiro, Tómas G Gunnarsson & José A Alves. Frontiers in Ecology & Evolution.

There is more about the information that is obtained from geolocators, how they work and the affects that they have on the individual birds that wear them in these two blogs:

blog roost flock


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

 

 

 

January to June 2019

blog CU postOne or two WaderTales blogs are published each month. The series is UK-based with a global reach. Suggestions of newly-published research on waders that might be of interest to birdwatchers who appreciate waders/shorebirds are welcomed. I am particularly keen to give feedback to colour-ring readers; they provide a huge amount of information that lies at the heart of these stories.

Click on a link in bold to read an individual blog.

You can sign up to receive an e-mail notification when a new blog is published.


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.

Managing water for waders

blog L sittingYour task, should you choose to accept it, is to turn farmland into a haven for breeding waders. The only tools you have at your disposal are tractors and cows and we will give you permission to pump water out of nearby rivers when conditions allow. That’s how it started. These days the diggers look big enough to use on a motorway construction site!

If your aim is to maximise the number of pairs of breeding waders on your lowland wet grassland farm or nature reserve, then one of the key issues is to get the water levels right. The first  part of this blog focuses on providing an appropriate mix of ditches, pools and grazed grassland for species such as Lapwing, Redshank and Snipe and then keeping everything wet enough (but not too wet) during the important chick-rearing season.

The second part of the blog is not just about maximising the number of breeding waders on a nature reserve. It’s also about working with neighbouring farmers, in order to secure fresh water supplies for the future and to reduce the risk of salt-water inundations, associated with sea level rise. In the long-term, stakeholder engagement has proved far more important than habitat management, as you will read below.

Understanding water levels

When developing lowland wet grasslands for waders, an extra five cm of late-winter rain can make a huge difference, especially if you can capture as much as possible of the rain that falls or can draw water from a swollen river. Mark Smart, the Senior Site Manager for the RSPB’s Berney Marshes and Breydon Water reserve in East Anglia, understands grazing marshes and how to capture and distribute water, in order to provide the muddy edges where wader chicks find insects. An aerial photograph of Berney Marsh shows how Mark has designed a special landscape to capture winter rain – one that is ideal for Lapwings and other waders.

Mark has taken the lessons he has learnt on RSPB nature reserves and shared them widely, a contribution to conservation that earned him one of the 2018 Marsh Awards for Wetland Conservation from the Wildfowl & Wetland Trust.

From wheat and barley to ducks and waders

blog cow eatingThe marshes of the Norfolk Broads have been drying out for 2000 years. By the 1970s, and after over 400 years of farming, the Halvergate Marsh complex was on the point of being fully drained and by 1985 much of the wet grassland in which waders formerly bred had already been lost and turned into arable fields. At this point, the RSPB made its first purchase of land, as they tried to retain at least some of the threatened habitat which is so important to winter wildfowl and summer waders. At the same time, campaigning by local and national conservationists secured legal protection for the unique Broadland scenery and the species that rely upon the habitats it contains, thereby halting the advance of the combine harvesters.

The importance of Halvergate Marshes for wildlife has long been known, and in 1987 it became the site of the UKs first Environmentally Sensitive Area – the prototype for subsequent agri-environment schemes. Lowland wet grasslands are traditionally drained using ‘footdrains’ – narrow, shallow channels that connect low-lying parts of the fields with surrounding ditches, in order to drain them.

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The muddy edges of a footdrain

These same footdrains can be used to hold and manage surface water levels within fields, by blocking the ditch connections with sluices. In the 1990s, Mark pioneered the design and deployment of footdrains on lowland wet grasslands, and the kit needed for their construction. Through his skills, enthusiasm and collaborations with grassland managers throughout lowland England, footdrains and the water that they can contain are now a common sight. Many generations of wader families have enjoyed the invertebrate food that footdrains support.

The breeding season for waders is very short – with the first Lapwing claiming territories in March and most chicks fledged by the end of July. Outside these months, these wet grasslands provide excellent grazing for geese and ducks in the winter and cattle in the summer. The task of nature reserve managers is to work with graziers to try to ensure that cattle deliver appropriate sward heights for winter wildfowl and summer waders.

Not just water

fence 2By creating a hot-spot for nesting birds, within an intensively-farmed landscape, land-managers also produce a food-rich area for predators, attracted in by concentrations of eggs, chicks and sitting adults. Restricting the activities of species such as foxes and crows is an important part of the role of an RSPB warden, carried out through site management and active control measures. By focusing these activities in the winter period, the RSPB’s Halvergate Marshes team are able to stop corvids and foxes from setting up territories within the area that is managed for breeding waders. Electric fencing can help to prevent foxes moving onto the site in spring, while changing the way that core wader areas are managed helps to reduce fox/nest interactions by, for instance:

  • Adding shallow ditches in the right places can break up the site into compartments and reduce the likelihood that nests will be predated.
  • Leaving areas with long grass, that is good for small mammals and the mustelids and foxes that prey upon them, can change the focus of hunting activities.
  • Erecting temporary fencing, during at least the early part of the nesting season, can both provide protection and potentially increase the synchronicity of nesting attempts and hence the ability of birds effectively to mob predators.

blog L chickThere is more about these measures in these blogs, with links to papers from the RSPB and University of East Anglia team of conservation researchers:

There is annual management of the Berney site too, with foot-drains to be re-cut, spoil to be spread in ways that can provide a mix of water-levels and more muddy edges, and rotovation of some areas to increase the diversity of habitats. These techniques might seem rather different to the ones that are used by farmers but many of the other operations at Berney are the same as would be seen outside nature reserves, with fences to mend, stock to manage and creeping thistle and rush to ‘weed-wipe’.

blog rotovation

Rotovation, carried out in dry conditions, adds heterogeneity and creates bare, muddy areas

Measuring success

The development of Berney Marshes has been hugely successful. Back in 1987, there were only 13 pairs of wader breeding on the site – nine pairs of Redshank and four of Lapwing. The total for 2019 looks like being about 270 pairs – over twenty times as many.

blog RK graphRedshank: The graph alongside illustrates how Redshank numbers have changed across the decades. At the same time as Breeding Bird Survey (BTO, JNCC & RSPB) for England results revealed huge declines, with a loss of nearly half in the period 1995-2017, Redshank pairs on Berney Marshes have been increasing. Even on this site, there is a suggestion that the peak number of pairs may be in the past. Given the pressures on breeding Redshank on saltmarsh habitats (blog: Redshank – the ‘warden of the marshes’), providing breeding habitat in coastal marshes, inside sea-walls, may be particularly important. Hopefully, the latest earthworks (see later) will create more space for Redshank.

Left image below shows Redshank nest in a clump of grass. The right image shows a Lapwing nest in a newly-rotovated patch. 

 

blog L graphLapwing: Between 1988 and 1998, the number of pairs of Lapwing rose from 14 to 79, reaching a peak of 157 in 2010. Numbers vary, according to spring weather and water levels, with between 83 and 130 pairs in the years 2011 to 2019. The national decline in England was 28% between 1995 and 2017 (Breeding Bird Survey) – not quite as drastic as for Redshank but still worrying. Intensive studies at Berney have shown that productivity is only high enough to boost numbers in some years. The latest project by UEA and RSPB conservation scientists involves trialling temporary electric fences to provide protection for first clutches and broods. Hopefully Berney can become a net exporter of Lapwings in most years.

Oystercatcher: There were no Oystercatchers breeding at Berney back in 1997. The peak number of pairs was 18 in 2009, with an average of ten pairs in subsequent years. Nationally, numbers in England have increased but with major declines in Scotland, which is the species’ heartland within the UK. This is discussed in an earlier WaderTales blog.

Avocet: The RSPB’s logo species has been hugely successful, nationally, with the help of protection and habitat creation. The first pair of Avocets bred at Berney in 1992 and pairs have bred in most years since then, with over thirty pairs in nine years but no pairs in 2013 or 2014. The 2019 count is 35 pairs and there is potential for further growth in numbers across the site, with the creation of more island homes (see below).

blog combine AV

Combined harvesters have been replaced by nesting Avocets

Snipe: Despite all of the excellent habitat creation work, there have never been more than 8 pairs of Snipe recorded on Berney and only between 0 and 3 pairs in each of the last ten years. The underlying soils at Berney are clay-based, which may not suit this species.

Sharing the knowledge

blog RK ringed

Redshank chick: science is an important feature of the work at Berney

Although it’s great that the RSPB has been able to buy and develop land for breeding waders in the Yare Valley, the impact of their work has been far larger, thanks to management agreements with other landowners. Mark Smart and the RSPB have set up Broads Land Management Services, to deliver wet grasslands that attract the top tier of conservation payments for farmers working in the Broads. Much of the recent work has been part of the Water Mills and Marshes Project, funded by HLF and led by the Broads Authority. Using specialist ditch-cutting and spoil-spreading equipment, the team has been able to create wet features within top-quality grazing fields. This is not just a local initiative; the kit and the advice have had impacts on farms and nature reserves across the country.

For his work for wetland conservation, Mark Smart received a Marsh Award for Wetland Conservation from the Wildfowl & Wetland Trust in 2018. “Mark Smart received his award for his 17 years managing RSPB Berney Marshes in the Norfolk Broads. Over this period, he brought together landowners, conservationists, local authorities and scientists to improve the marshes for wildlife. Today more than 300 pairs of wading birds nest there each spring, and more than 100,000 waterbirds return to it each winter.”

blog somerleyton

Illustrations above shows work that has been completed at Somerleyton in Suffolk and a newly-fledged Lapwing.

blog DutchWorking with his wife, Jen Smart, who is a Principal Conservation Scientist at the RSPB’s centre for Conservation, Mark has added a Dutch dimension to the RSPB’s advice work by co-authoring Meadowbirds on the horizon of southwest Friesland.  This report has just been published by the International Wader Study Group.

Climate – ‘the new normal’

Fresh water is an increasingly important commodity in East Anglia – for farmers and for nature reserve wardens, looking to maximise agricultural and wader chick production. More extreme weather patterns are already producing periods of drought and intense periods of rain, while a rising sea-level is increasing the salinity of rivers and limiting extraction opportunities. Broadland farmers are looking for a reliable water supply, the Environment Agency is looking for ways to reinforce sea defences and for places to store fresh water, in order to avoid flooding, and the RSPB wants to hold more water in the late winter that can be used to keep areas wet in the early summer. With some lateral thinking, many of the needs of these key stakeholders can be met in partnership projects, as shown below

The Environment Agency’s need for material to raise sea defences provided Mark Smart with an opportunity to provide more pools and scrapes for breeding waders. It was a win-win solution; free habitat creation work for the RSPB and minimal movement of the clay and top-soil that the Environment Agency needed. In the images below, you can see this work in progress and the islands that are now being used by nesting Avocets.

blog defences

The most recent project is an ambitious water storage and flood reduction scheme for the whole of Halvergate Marsh. This will keep salt water out of these important grazing marshes and store fresh water for summer use. The £2 million Halvergate Marshes Water Level Management Improvement Scheme is a joint initiative, funded by DEFRA and delivered by the Water Management Alliance. The project involves a large number of stakeholders, including the Broads Internal Drainage Board, RSPB and neighbouring farming estates.

blog 8 km

The Water Level Management Improvement Scheme is a huge undertaking, with 8 km of new ditches, 240 piped culverts and 12 big sluices, that will create storage for 60,000 cubic metres of fresh water and systems to distribute this water over the course of a dry East Anglian summer. One of the most impressive features of the project, illustrating the imagination of the design team, is the Higher Level Carrier, a ‘flyover’ ditch system that passes over the top of existing wet grazing land to get water to some of the driest part of Halvergate Marshes (left picture below). This high-level water transportation route was constructed using locally-sourced clay, thereby creating shallow pools around which yet more waders are already nesting.

blog overpass

When designing this project, the opportunity was taken to develop opportunities for birdwatchers to see the birds that will be drawn into the wettest areas, by making sure that the ‘best bits’ are close to public access points on the Weavers’ Way, the 61 mile (100 km) long-distance path running from Cromer to Great Yarmouth.

Aspirations

Mark Smart has not finished yet! Plans are afoot to develop the RSPB’s land that is closest to Great Yarmouth, recently purchased using a WREN grant. If agreed, this can provide an alternative, safe high-tide refuge area for tens of thousands of waders and wildfowl that roost on the mud and saltmarsh at the mouth of the Yare. Their current high-tide refuge is threatened by sea-level rise and developments proposed for the outskirts of Great Yarmouth.

blog aspirations

This proposed roosting area will be part of an extension to the Halvergate Marshes Water Level Management Improvement Scheme, which will add another 10,000 cubic metres of water storage. Alongside flood alleviation and fresh-water conservation, this scheme will create fifty hectares of additional shallow wader and waterfowl scrapes adjacent to Breydon Water.

blog Wood spThe new scrapes should not only attract wintering and breeding birds but also many passage waders, such as the Wood Sandpiper pictured to the right. The whole scheme has the potential to be another win-win-win-win, for the owners of low-lying properties, for Broadland farmers, for internationally important bird populations and for local and visiting birdwatchers.

Read more

Information about the RSPB’s Berney Marshes & Breydon Water reserve can be found on the RSPB’s website. Click here

There is information about the Water, Mills and Marshes project here.

blog AV


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

 

Chicks and ticks

How do ticks affect Golden Plover chicks? pic chick on moveBy utilising data from an existing study, David Douglas and James Pearce-Higgins have discovered that Golden Plover chicks that carry more sheep-ticks Ixodes ricinus have a lower chance of survival. Their findings are written up as a paper in Bird Study. The work is only based on a small sample and the data don’t identify the mechanism that leads to increased mortality but, given the current interest in the biological effects of ticks, the findings are interesting.

Costs of carrying ticks

Carrying ticks has three potential effects on wader chicks

  • Ticks suck blood, which could be costly.
  • Ticks can introduce diseases, via tick-borne bacteria and viruses.
  • There may be effects on feeding efficiency, via impaired vision, hearing etc.

Sheep ticks act as vectors for a variety of pathogens, including the louping ill virus (LIV) which can affect a range of domestic and wild mammals, as well as wild birds. LIV is known to cause high mortality of Red Grouse chicks but there has been no previous assessment of the effects of sheep ticks on other moorland birds, such as breeding waders. It should be noted that wader chicks eat ticks – so they are not ‘all bad’.

pic ticks

Ticks can clearly be seen on the unfeathered lower eyelid of this young chick. There are smaller ticks attached to the gape at the base of the beak as well.

All sorts of things could influence the probability of tick infestation in birds:

  • Ticks may be able to survive better in warmer conditions.
  • Tick numbers can be affected by the number of mammalian hosts.
  • Mammal to bird transfer could be affected by land management and habitat structure.

Ecological interactions between ectoparasitic ticks and waders are not well understood. Given possible increases in tick abundance with climate change, the authors of the new study felt that it would be useful to test whether ticks have detectable effects on the Golden Plover chicks that carry them.

Spotting an opportunity

The Golden Plover chicks that provided the data used in this paper were caught as part of a wind farm study at Gordonbush in northern Scotland, a site made up of 33 km2 of blanket bog. There is more information about the study in these two papers:

pic habitat

Gordonbush, prior to the development of a wind farm

Tick numbers on Golden Plover chicks were collected at the time of ringing (within 24 hours of hatching) and during subsequent recaptures. Recapture was facilitated by locating tagged birds using radio-location. On each capture, chicks were weighed and the number of ticks visible on the bare parts of the head (around the eyes and bill) were counted. Most ticks attach themselves to the bare parts of the head and neck.

Variation in tick loads on Golden Plover chicks

pic nesting ad

Incubating adult. Golden Plovers commence primary moult at the start of (or before) incubation. Read more here.

The number of sheep-ticks found on these Golden Plover chicks was higher than those previously reported for waders but were within the range of those found on Red Grouse on moorland. Previous wader studies had been focused upon areas with sheep, which were routinely treated to reduce tick infestations. In the current study it was found that:

  • 90% of chicks were carrying ticks, with between 1 and 12 ticks being found on each affected Golden Plover chick. The highest tick-load was found in mid-age chicks.
  • Tick loads were higher during periods with warmer maximum temperatures and when chicks were estimated to have moved through taller vegetation between recaptures.
  • Chick growth rates were depressed by high tick-loads, especially when temperatures were warmer.
  • Of the 21 chicks, 4 fledged, 13 died and the outcomes for the other 4 were unknown. Half of the deaths appeared to have been due to predation and half to starvation/exposure.
  • Chicks that were heavier (for their ages) were more likely to survive. Those with higher tick loads (for their ages) were less likely to survive.

With the small sample size, it was not possible to detect a correlation between tick load and chick growth rates but low survival was correlated with high tick-loads. This had not previously been documented for waders.

Implications for wader conservation

pic red deerGordonbush is an area where there is no grazing by domestic animals so the likely mammalian tick-hosts are Red Deer in particular and also Mountain Hare. The correlation between warmer weather and tick numbers, found in this study, could be explained by increased tick activity, while the link to taller vegetation may well be explained by ticks seeking damper microhabitats. In their discussion of the results, the authors suggest potential ways that ticks and waders, of different ages, might interact. Anyone looking to expand the work, in order to understand the mechanics of tick infestation, is likely to spend more time looking at ticks than waders!

pic Curlew

Could ticks be reducing survival probabilities for  young Curlew?

The authors of this paper were not able to test for the presence of disease (such as LIV) in their Golden Plover population but this is a plausible cause of the increased probability of mortality. In a study in Yorkshire by Newborn et al. (2009), no evidence was found of LIV in wader chicks, whereas it was present in 3.6% of a sample of Red Grouse chicks at the same sites. Newborn and colleagues report that a single Eurasian Curlew chick has previously been recorded to be seropositive for LIV. In the Newborn study, the lowest incidence of ticks among waders was in Lapwings (6% of broods), followed by Golden Plover (47% of broods had ticks) and Curlew (91% of broods). There is a hint, in these data, that ticks might more commonly attach themselves to wader chicks that are found in taller vegetation.

Despite high tick loads on chicks, and the correlation with lower chick survival, the overall percentage of Golden Plover chicks known to fledge in the Gordonbush study (19%) is comparable with other studies. Perhaps ticks are only causing the deaths of chicks that would have died anyway?

pic Red GrouseThe authors suggest that no case should be made for tick-control, to help breeding waders, until it is clear whether tick-based chick mortality limits Golden Plover and other wader populations on moorland. In the paper, they argue that previous attempts to reduce moorland tick abundance and tick loads on Red Grouse have failed to detect convincing evidence of improvements in grouse survival, breeding success or post-breeding densities.

A range of methods have been deployed in attempts to reduce moorland tick abundance and tick loads on Red Grouse, including reducing densities of mammalian tick hosts (Mountain Hare and deer) and deploying acaricides (as used in sheep dips). See this link to material from GWCT on ticks and Red Grouse.

In conclusion

pic older chickWork at a single site over two years appears to have documented a level of tick infestation in Golden Plovers that is associated with chick mortality. It is not clear how chicks are being affected, particularly given that there is insufficient evidence thus far that ticks affect chick growth rate.

The authors collected the data analysed in this paper for other studies – the focus was not on tick effects – and they hope that funding might be found for future research focusing upon the associations between ticks and waders, other birds and other animals. Until that happens, it would be useful if shorebird biologists who repeatedly handle wader chicks, in order to measure growth and survival rates, could routinely record the presence or absence of ticks.

Given that warming temperatures could lead to increased tick abundance, this seems to be a good time to discover more about tick behaviour, the importance of ticks as a food source for wader chicks and whether tick-loads are reducing growth rates and fledging success in other wader species.

Paper

This research is published in the BTO journal Bird Study. Click on the details below to link to the full paper:

Variation in ectoparasitic sheep tick Ixodes ricinus infestation on European Golden Plover chicks Pluvialis apricaria and implications for growth and survival. David J. T. Douglas and James W. Pearce-Higgins. Bird Study. June 2019.

pic bottom


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