There is nothing more obvious than an Oystercatcher sitting on his or her nest, but a brooding Snipe can be invisible until almost trodden upon. Which strategy works better: nesting in plain view but laying cryptically camouflaged eggs or hiding yourself and your nest in a clump of grass? Which species is most likely to hatch a successful brood of chicks and in what circumstances? In a 2020 paper in IBIS, Becky Laidlaw and colleagues analysed nest site characteristics and nest locations of 469 wader nests in Iceland in order to provide some answers
The perils of ground-nesting
Almost all waders are ground-nesters, which makes them highly vulnerable to a wide range of nest predators. To reduce the risks of predation, different strategies have evolved. In some species, nests are placed out in the open, and the camouflage is provided only by mottled egg colouration that resembles the background. In other species, nests are secreted in vegetation, meaning eggs and incubating adults are concealed from predators.
In both groups of species, the risk of nests being predated might vary, depending on the surrounding habitat. For open-nesting species, for example, clutches that are laid in large patches of similar habitat may be harder for predators to locate. The same could apply to closed-nest species that hide their nests; Snipe nests may be tricky to find in extensive areas of long grass but perhaps more at risk if there are only a few suitable clumps of long grass that predators need to check out.
Iceland: a wader factory
As discussed in previous WaderTales blogs, particularly Do Iceland’s farmers care about wader conservation? Iceland is hugely important as a European ‘wader factory’. As farmland elsewhere has become less suitable for species such as Redshank and Snipe, the global importance of the country has increased (see table alongside for most recent figures from an AEWA report) With this in mind, it is important to understand the factors that underpin the population dynamics of Iceland’s breeding waders.
Working in South Iceland, Becky Laidlaw and her co-authors tried to find as many nests as possible during the summers of 2015 and 2016. This area is largely a mosaic of open habitats, although there are more patches of forestry than there were twenty years ago. Most of the Southern Lowlands area is farmed, on a gradient between intensive and semi-natural, and this is reflected in the distribution of breeding waders (see Farming for waders in Iceland).
For this project, nests were located by surveys from vehicles and on foot, through observation of incubating adults, systematic searching, incidental flushing of incubating adults and rope-dragging (dragging a 25 m rope, held between two fieldworkers, lightly across vegetation) to flush incubating adults.
The analysis in the resulting paper in IBIS focuses on 469 nests of three open-nesting species (Oystercatcher, Golden Plover and Whimbrel) and three species that hide their nests in tall vegetation (Redshank, Snipe and Black-tailed Godwit). The team recorded the habitat and vegetation structure around each nest (at the nest, within a 5 m x 5 m square and in a wider 50 m x 50 m square) and worked out which nests hatched successfully and which were predated. The date and time of predation were determined, where possible, with nest-cameras providing extra information for some nests. Cameras captured nest-predation events involving Arctic foxes, Arctic Skuas, Ravens and sheep.
Interestingly, 2015 and 2016 were very different wader breeding seasons. The graphic below shows the mean temperatures for the months from April through to July (encompassing the wader breeding season at this latitude) were much cooler in 2015 than in 2016, representing average monthly difference of between 1.5°C and 2.5°C. At high latitudes these figures translate into very different rates of vegetation growth.
First, find your nest
When nests were first located, their positions were marked and referenced using GPS. Eggs were floated in water to provide an estimate of laying date and thereby predict hatching date. As the chick develops within an egg, the density of the egg falls. A newly laid egg will lie on the bottom of the flotation vessel. Over the next few days the ‘blunt end’ rises until the egg is still touching the bottom but vertical. Eggs in the late-development stage float ‘point-end-down’, with the latest eggs floating at an angle to the vertical (method described by Liebezeit et al.).
Nests were considered successful if one or more eggs hatched, and predated nests were defined as those that were empty in advance of the predicted hatch date or those without any eggshell fragments in the nest (a sign of successful hatching). To determine the time and date of nest failures, iButton dataloggers were placed in a randomly selected subsample of nests. These loggers recorded a temperature trace every ten minutes. A sharp and permanent decline in nest temperature below incubation temperature indicates nest predation. In both study years, motion-triggered cameras were deployed on a sample of open-nesting species to determine the predator species active on these nests.
When each nest was first located, the percentage of eggs visible from directly above the nest was estimated and the habitat surrounding each nest was assessed in the field at three spatial scales: the nest cup, the 5 m x 5 m and the 50 m x 50 m area surrounding each nest. Details are in the paper.
Which nests survive through to hatching?
Over the breeding seasons of 2015 and 2016, the outcomes of 469 wader nests were assessed. 259 hatched successfully (55%), 192 were predated (41%), 13 were abandoned, 7 were trampled and 2 were mown. A nest-loss rate of 40% is fairly typical for ground-nesting waders, when compared to studies in different countries and habitats.
Daily nest predation rates did not vary significantly in relation to the habitat heterogeneity or the extent to which the dominant habitat covered the area surrounding the nest, at either 5 m x 5 m or 50 m x 50 m scales. Most clutches were laid in habitats that were the same or similar to the surrounding areas. Where there were differences, the dissimilarity between the habitat at the nest cup and in the surrounding area did not influence daily nest predation rates for open- or closed-nest species. Although nest predation is high, at about 40%, incidence of predation events appears to be unpredictable – or even random.
Daily nest predation rates were significantly higher for closed nests (Redshank, Snipe and Black-tailed Godwit nests) in which a greater percentage of the clutch was visible. This suggests that the onset and rate of vegetation growth could potentially constrain the availability of suitable nesting locations for these species, and hence influence nest success, particularly among early season nests. This has been studied in Icelandic Black-tailed Godwits by José Alves and colleagues and is described in From local warming to range expansion.
For closed-nest species, the visibility of nests was significantly greater during the early part of the 2015 breeding season, when compared to 2016, due to slower grass growth in cooler conditions. The higher predation rate of more visible nests of closed-nesting species was apparent even though nests were predated up to three weeks after egg visibility was measured. These findings suggest that early nesting attempts by concealed-nest species are unlikely to be successful in years when vegetation growth is delayed or slow. There can be major benefits of hatching early, with recruitment into breeding populations typically being lower for later-hatched chicks, so vegetation growth rates are likely to be really important to species that conceal their nests (Redshank, Snipe & Black-tailed Godwit in this study). However, given the ongoing trend for warmer springs at subarctic latitudes, the conditions in which early nests can only be poorly concealed are likely to be reducing in frequency.
Perhaps surprisingly, nest predation rates were similar for open-nest and concealed-nest species and did not vary with vegetation structure in the surrounding landscape. However, nest-concealing species were about 10% more likely to have nests predated when the nests were poorly concealed, and the frequency of poorly concealed nests was higher at the start of the breeding season in colder conditions.
The paper at the heart of this blog is:
Vegetation structure influences predation rates of early nests in subarctic breeding waders. Rebecca A. Laidlaw, Tómas G. Gunnarsson, Verónica Méndez, Camilo Carneiro, Böðvar Þórisson, Adam Wentworth, Jennifer A. Gill and José A. Alves. IBIS. doi:10.1111/ibi.12827
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.