In most northern waders, post-breeding moult is a distinct phase – between autumn migration and the start of winter – but it’s different for Lapwings
Primary moult is one of the key stages of a wader’s life. Knowing the length of the period in which birds are constrained by this energetic process may be really important to our understanding of the whole annual cycle of waders. In Oystercatchers, for instance, it has been shown that they cannot finish their primary moult in periods of low food abundance. One or more of the outer pairs of primaries are retained, becoming more faded and worn as the next twelve months pass and providing a signal of a previous period of stress. [See BTO report 238 by Atkinson et al.]
Lapwing moult was first studied in detail in the 1970s using an innovative technique that did not require birds to be captured. By collecting primary feathers dropped by a flock of Lapwings that spent several weeks near their Buckinghamshire home, David and Barbara Snow were able to estimate the duration of moult. They did not start picking up feathers until August in 1974 but they were ready for the start of the 1975 season and captured a whole season’s worth of data that year, visiting the site every four days during a long, dry summer.
Primary wing moult
For flocks of waders in Britain and Ireland, autumn moult is generally squeezed in between their return from the arctic and sub-arctic breeding grounds and the start of winter, when days get shorter, the weather gets colder and much of the prey becomes less numerous and harder to access. In the late summer, adult birds start to discard the colourful feathers that they grew in spring and the main wing and tail feathers that have served them well for the best part of a year. The key period of moult is the time in which the ten primary feathers are replaced. The inner one (primary 1) on each wing is dropped first and the outer one is last. Each of the ten primary feathers is scored from 0 to 5, so an old wing has a score of 0 and a new wing has a score of 50. During the process, there is a gap in the wing, where new feathers are growing. A typical bird in mid-moult might have ten feathers scored as 5554332100 (moult score 28), with primary four nearly fully grown and primary eight newly discarded. Unsurprisingly, flight capacity is impeded. It’s not a great time to migrate – unless you happen to be a Lapwing.
David & Barbara Snow
David & Barbara Snow were each respected ornithologists in their own right and David was the author of the BTO’s Guide to Moult in Birds and senior scientist in the bird section of what is now the Natural History Museum. Wherever they lived or worked – from Trinidad to Tring – they observed the birds around them, so when a flock of Lapwings gathered to moult near their home they decided to collect the feathers that they dropped. They discovered that it is possible to use the individual patterns of white spots and the lengths of primary feathers in order to assign primary number to discarded Lapwing feathers, and that this could be done with certainty for primaries 1, 2, 7 and 10 (with 1 being the innermost). Reference to a small sample of BTO moult cards allowed David & Barbara to assign moult scores of 1, 4, 29 and 41 to the points at which these four distinct feathers had been dropped and collected. The synchrony of moult in the two wings was evident: “Several times we found pairs of feathers lying exactly where they had been dropped, separated by a body width”.
Much of what the Snows discovered from recording in detail the feathers shed by their local flock of 70 and 120 moulting Lapwing in 1974 and 1975 is still relevant. They suggest that the collection of dropped feathers might be a useful adjunct to the study of moult in situations where birds roost in the same place on a daily basis, as might be the case for flocks of gulls or individual raptors.
David and Barbara also collected secondary feathers and tail feathers. As there is a difference between the outer tail feathers of adult and first-year birds they were able to show that early moulters included a higher proportion of first-year birds. The paper is well worth a read. David W. Snow & Barbara Snow (1976) Post-breeding Moult of the Lapwing, Bird Study, 23:2, 117-120
As far as I can ascertain, nobody has tried to use this technique again. I feel partially responsible, having written a follow-up paper with Clive Minton that pointed out some flaws in the methodology. These related to a lack of linearity in the progression of primary moult, as measured using the normal score of 0 to 50 based on 0-5 for the ten individual feathers, and the time taken to complete moult once the last primary has been dropped. Using data from Lapwing caught in cannon-net catches in the English Midlands, Clive Minton and I found that Lapwing moult starts very early, with median commencement and completion dates of about 7 June and 29 September – an estimated duration of 114 days and much longer than that suggested in the figure produced by the Snows. G. F. Appleton & C. D. T. Minton (1978) The primary moult of the Lapwing, Bird Study, 25:4, 253-256.
Lapwings are strange
Most waders that we see in the UK have distinct moult and migration phases to the post-breeding portion of their annual cycle. After having successfully raised a brood of youngsters or attempted so to do, Turnstones, Bar-tailed Godwits, Knot etc. leave their nesting areas in the high Arctic and fly thousands of kilometres, arriving on our shores in July and August. Although some populations, such as most Dunlin of the schinzii race, only pause to fatten up and then move on to moult further south in northern Africa, many waders that use the UK during autumn undergo a full moult. One can imagine circumstances in which a small flock of inland-moulting Curlew that use the same roost site on a daily basis might be studied in the same way as the Snows studied their Buckinghamshire Lapwings, as long as the lengths and patterns on individual feathers allow the primary number of some of the individual primaries to be assigned with a degree of certainty.
Unlike most of their wader cousins, many Lapwings migrate while moulting their primary feathers. Arctic species have to make long-hop migrations and are restricted in their choice of feeding areas by the distribution of estuaries but Lapwings make short movements and can break their journeys as often as they like. The North Sea is the biggest barrier they face when making journeys from far east in Russia but a crossing from the Netherlands to Norfolk is less than 150 km. Lapwings arriving on the east coast of England have been seen to have gaps in their primaries, despite the fact that these feathers deliver most of the lift and propulsion.
Moult strategy is not the only strange thing about Lapwings; they have also been shown to exhibit abmigration, as described by Chris Mead in a paper with Jim Flegg and Chris Cox in 1968. Although most British-bred Lapwings return to their natal areas to breed, as shown by Pat Thompson et al in 1994, a small number of birds breed a long way from where they were hatched, including Russia, Norway and Sweden. The birds that go on to breed in continental Europe are assumed to join flocks which happen to be travelling east in spring. Given that British-hatched birds as old as 12 years of age have been found breeding in Russia, it is assumed that, having once travelled to a new breeding area**, Lapwings will repeat the journey each year. Genetic mixing at large geographic scales would potentially explain a lack of morphological variation across the large range of the species.
** In the summer of 2022, two female Lapwing that were satellite-tagged in the Netherlands amazed researchers when, having lost their first clutches, they flew east to Russia and Ukraine, presumably for a second nesting attempt. 2022 was a particularly dry year and it is possible that these two birds went in search of better conditions, joining flocks of Lapwing that were heading to the coldest part of the breeding range. There’s more (in Dutch) on the SOVON website.
Another possible – but unlikely – explanation is that the researchers happened to tag two Lapwings that did not ‘own’ the nests upon which they were caught. Nest cameras have taught us that the brooding instinct is very strong. For instance, during a predation experiment that involved creating dummy nests, Becky Laidlaw (University of East Anglia) witnessed Lapwings trying to brood her quail eggs.
Collect some feathers
Moult is currently only studied when birds have been caught but perhaps it might be possible to use digital photography to establish the primary moult duration of colour-ringed individuals or to use the Snows’ technique of collecting feathers from a moulting flock of birds over a period of a few weeks. On an annual basis there may even be opportunities to monitor change? A reduced rate of moult, as measured through the collection of discarded primaries, could provide a way to assess inter-annual variation in stress. I wonder if moult periods have increased for birds on the East-Asia Australia Flyway, where they are experiencing reductions in feeding opportunities on their southward migration via the Yellow Sea? Wouldn’t it be interesting if a technique first described by the Snows forty years ago could be used by a new generation of wader biologists when investigating the latest issues of conservation concern?
The following 1980 paper has been brought to my attention, in which Arend Jan van Dijk wrote about the moult of black-tailed godwits, based upon visual observations of the gaps in the wings of overflying birds and the collection of shed feathers: Observations on the moult of the Black-tailed Godwit LIMOSA 53 (2): 49-57.
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.