The effects of raptor predation on wintering wader populations at the Tyningharne estuary, southeast Scotland

Raptor predation on waders was studied by direct observation of raptors hunting a known wader population and subsequent recovery of dead waders. In each of three winters, raptor predation was shown to be the most significant cause of mortality in most small wader species, Sparrowhawks Accipiter nisus , Merlins Falco columbarius and Peregrines F. pere-grinus attacked waders with a success rate of 11.6%. 8.8% and 6.8%, respectively. Most waders attacked or found dead were Redshank Tringa totanus and Dunlin Calidris alpina; most were killed by Sparrowhawks. Kleptoparasitism of raptors carrying prey by Carrion Crows Corvus corone significantly increased the winter mortality of some waders. Redshank populations were most affected by raptor predation: over 50% of the total population (which was found to be closed during most of the winter) and over 90% of the juvenile population were taken in two winters: juvenites were more likely to be killed by raptors.     

Regional trends in coastal wintering waders in Britain

Regional trends in the populations of waders wintering largely on estuaries were analysed using data collected by the Wetland Bird Survey between the winters of 1969/70 and 1995/96 to assess whether trends seen at the national level were repeated across regions of Britain. Not all were. Over this period, the numbers of seven of the 12 species studied increased in south and southeast England, but remained static or declined in southwest England and south Wales. Possible explanations for this are proposed. Increases in the number of Curlews Numenius arquata in the north and northeast of Scotland occurred after 1981/82, when the species stopped being a legal quarry species. A rapid decline in Oystercatcher Haematopus ostralegus and Knot Calidris canutus numbers in east England after 1988/89 may be linked to a change in the Wash shell-fisheries. Warmer winters may be making it less essential for birds to winter in the milder west of Britain. In two pairs of regions, northeast and southeast Scotland, and in northwest England/north Wales and northeast England, waders showed similar long-term population changes. The long-term population trends of Oystercatchers in the different regions correlate to those of Ringed Plovers Charadrius hiaticula and Knot, while those of Ringed Plovers are very similar to those of Bar-tailed Godwits Limosa lapponica. In some regions of Britain, the population trends of some species varied a lot between sites, a factor that is an important consideration when determining the size and geographical position of regions for future regional indexing. The present separation of regions based on water catchments and geography has given a useful insight into population changes at a finer spatial scale but it could be improved further with added biological input.     

Vegetation structure influences predation rates of early nests in subarctic breeding waders

Ground-nesting species are vulnerable to a wide range of predators and often experience very high levels of nest predation. Strategies to reduce nest vulnerability can include concealing nests in vegetation and/or nesting in locations in which nests and eggs are camouflaged and less easy for predators to locate. These strategies could have important implications for the distribution of ground-nesting species and the success rates of nests in areas with differing vegetation structure. However, the factors influencing the success of nest concealment and camouflage strategies in ground-nesting species are complex. Here we explore the effects of local vegetation structure and extent of nest concealment on nest predation rates in a range of ground-nesting, sympatric wader species with differing nest concealment strategies (open-nest species: Oystercatcher Haematopus ostralegus, Golden Plover Pluvialis apricaria and Whimbrel Numenius phaeopus; concealed-nest species: Black-tailed Godwit Limosa limosa, Redshank Tringa totanus and Snipe Gallinago gallinago) in south Iceland, in landscapes that comprise substantial variability in vegetation structure at a range of scales. We monitored 469 nests of these six wader species in 2015 and 2016 and ~40% of these nests were predated. Nest predation rates were similar for open-nest and concealed-nest species and did not vary with vegetation structure in the surrounding landscape, but nest-concealing species were ~10% more likely to have nests predated when they were poorly concealed, and the frequency of poorly concealed nests was higher in colder conditions at the start of the breeding season. For concealed-nest species, the reduced capacity to hide nests in colder conditions is likely to reflect low rates of vegetation growth in such conditions. The ongoing trend for warmer springs at subarctic latitudes could result in more rapid vegetation growth, with consequent increases in the success rates of early nests of concealed-nest species. Temperature-related effects on nest concealment from predators could thus be an important mechanism through which climate change affecting vegetation could have population-level impacts on breeding birds at higher latitudes.     

An assemblage of wintering waders in coastal Kenya: activity budget and habitat use

To contribute to the poorly known ecology of Palearctic immigrant waders in East Africa, we studied the foraging system (roosting and foraging behaviours, 24‐h activity rhythm, time‐activity budget and habitat use) of eleven wader species overwintering on the Kenyan coast. The waders congregated in communal roosts and completely overlapped in their foraging rhythms, which in this tidal environment were completely determined by tides, and which were independent of the dark/light cycle. All species, both tactile and visual foragers, fed both by day and by night. This study, along with similar findings for other areas of the Palaeotropics, shows that wader activity throughout the 24‐h day is essentially uniform, and is only slightly influenced by the dark/light cycle, but is modified by tidal levels. The total time devoted to feeding by each wader species was related to their body mass. Each species used a specific micro‐habitat for foraging.     

Variability in the area,energy and time costs of wintering waders responding to disturbance

Birds’ responses to human disturbance are interesting due to their similarities to anti‐predator behaviour, and understanding this behaviour has practical applications for conservation management by informing measures such as buffer zones to protect priority species. To understand better the costs of disturbance and whether it will impact on population size, studies should quantify time‐related responses as well as the more commonly reported flight initiation distance (FID). Using waders wintering on an estuarine area, we experimentally disturbed foraging birds on the Wash Embayment, UK, by walking towards them and recording their responses (FID, alert time, time spent in flight, time taken to resume feeding, and total feeding time lost). We present data for 10 species of conservation concern: Curlew Numenius arquata, Oystercatcher Haematopus ostralegus, Bar‐tailed Godwit Limosa lapponica, Grey Plover Pluvialis squatarola, Redshank Tringa totanus, Knot Calidris canutus, Turnstone Arenaria interpres, Ringed Plover Charadrius hiaticula, Sanderling Calidris alba and Dunlin Calidris alpina. Larger species responded more strongly, response magnitude was greater under milder environmental conditions, and responses varied over both small and large spatial scales. The energetic costs of individual responses, however, were low relative to daily requirements and disturbance events were unlikely to be frequent enough to seriously limit foraging time. We suggest, therefore, that wintering wader populations on the Wash are not currently significantly negatively impacted by human disturbance during the intertidal foraging period. This is also likely to be the case at other estuarine sites with comparable access levels, visitor patterns, invertebrate food availability and environmental conditions.     

The influence of landscape features on nest predation rates of grassland‐breeding waders

In Europe, lowland wet grasslands have become increasingly fragmented, and populations of waders in these fragments are subject to unsustainably high levels of nest predation. Patches of taller vegetation in these landscapes can support small mammals, which are the main source of prey for many predators. Providing such patches of habitat could potentially reduce levels of nest predation if predators preferentially target small mammals. However, predator attraction to patches of taller vegetation for foraging, shelter, perching and/or nesting could also result in local increases in predation rates, as a consequence of increased predator densities or spill‐over foraging into the surrounding area. Here we assess the influence of taller vegetation on wader nest predation rates, and the feasibility of managing vegetation structure to alter predator impacts. Between 2005 and 2011, the nest distribution and hatching success of Northern Lapwings Vanellus vanellus, which nest in the open, and Common Redshanks Tringa totanus, which conceal their nests in vegetation, were measured on a 487‐ha area of wet grassland in eastern England that is primarily managed for breeding waders. Predation rates of Lapwing nests increased significantly with distance from patches of taller vegetation, and decreased with increasing area of taller vegetation within 1 km of the nest, whereas neither variable influenced Redshank nest predation probability. These findings suggest that the distribution and activity of nest predators in lowland wet grassland landscapes may be influenced by the presence and distribution of areas of taller vegetation. For Lapwings at least, there may therefore be scope for landscape‐scale management of vegetation structure to influence levels of predation in these habitats.     

Short-term effects of reclamation of part of Seal Sands,Teesmouth, on wintering waders and Shelduck

Summary The invertebrate macrofauna of Seal Sands, Teesmouth, is very limited in species composition. Nereis diversicolor has a two-year life cycle; the larger size-class provides the main prey of the birds Pluvialis squatarola, Numenius arquata and Limosa lapponica. Hydrobia ulvae is an important food of P. squatarola and Calidris canutus. Small Carcinus maenas occur in late autumn and are taken by the larger shorebirds. Small Macoma balthica are also taken, but are scarce and not an important bird food. Tadorna tadorna and Calidris alpina subsist chiefly on species of small oligochaetes and polychaetes which occur at very high densities (Gray 1976). Feeding areas of the shorebird species show some segregation, particularly in groups of species taking the same prey. It is calculated that birds removed about 90% of the standing crops of large Hydrobia and Nereis during a single winter. This followed the reclamation of more than half the intertidal land used as feeding grounds by the birds.     

Low tide distribution of wintering waders and shelduck on the Severn Estuary in relation to the proposed tidal barrage

Waders and shelduck were counted at low tide on 162 sectors comprising 85% of the intertidal area (21 467 ha) of the Severn Estuary on 12 occasions during winter 1987/88. On average, 50% of birds present at low tide utilized just 13 sectors (12% of the area); 90% of birds occurred on only 56 sectors, leaving large expanses of intertidal sand virtually devoid of birdlife. Dunlin, the numerically dominant species, occurred widely on the middle and outer estuary, whereas shelduck predominantly occurred on the outer estuary and redshank around many tributary river mouths. Curlew, the most ubiquitous species, was the only one concentrated on the inner estuary. Severe gales in both late December and mid-January concentrated all main species within fewer sectors, probably by the short-term removal of surface sediment from substantial areas. It is estimated that the proposed tidal barrage would eliminate intertidal areas accounting for between c. 40% (for shelduck and curlew) and 80% (for redshank) of current total low tide usage by the internationally important populations present.     

Habitat selection,reproduction and predation of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.     

Artificial nest and seed predation experiments on tropical southeast Asian islands

Southeast Asia is rapidly losing native habitats and the consequences of this are poorly understood. Because habitat loss and disturbance can affect avian and seed survivorship, we conducted artificial nest and seed predation experiments on tropical southeast Asian islands. Data among islands and fragments or different forest types (e.g. primary versus exotic forest) within the islands are compared. On Singapore Island, predation among different forest types (primary, secondary and woodland) did not differ. Only at one of the sites, nest predation was higher at 75 m from the forest edge than at 25 m. In other sites, predation did not differ in relation to the distance from the forest edge. Predation among 10 small (0.8–1026 ha) Singaporean islands differed. However, none of the environmental variables (e.g. island area) could explain the predation differences. The lowest predation of both nests and seeds was recorded in the primary forest areas of a contiguous forest (25 500 ha) in central Java (Linggoasri). Small mammals were the main predators on Singapore and other surrounding islands. However, the index of potential predator abundance, overall, did not correlate with predation. While larger and more pristine forests may be better for avian and seed survivorship, pinpointing variables affecting both artificial nest and seed predation may be difficult.     

High water roost selection by waders: maximizing feeding opportunities or avoiding predation?

During the highest spring tides the intertidal sediment flats of estuaries are fully inundated at high water, and waders have no choice but to move to supratidal roosts, e.g. on open farmland, saltpans or beaches. However, in many estuaries during the lowest neap or intermediate tides there are sectors of upper intertidal sediment flats that remain exposed even at the peak of high water, and so waders have the choice of roosting either there or in supratidal sites. In the Tagus Estuary, Portugal, as elsewhere, waders use both types of roosts during high water. Our main objective was to understand what makes waders opt for one of these two types of available roosts. We monitored wader use of saltpans and intertidal roosts from spring to neap tides, and measured foraging and alarm behaviour, prey availability and disturbance by predators. Most of the wader species studied chose intertidal (mudflat) roosts whenever these were available, and only roosted in saltpans during the peak of spring tides. We hypothesized that this preference was explained either by an attempt (i) to continue feeding into the high water period, or (ii) to minimize predation risk. Extending feeding time into the high water period did not seem to be very relevant for roost choice because both prey availability and foraging activity were low in both types of roosts. However, predator disturbance was several times higher in the saltpans than in the intertidal roosts, suggesting that this factor may be the determinant in the choice of roost type.     

Effects of microhabitat on palm seed predation in two forest fragments in southeast Brazil

The establishment of plants depends crucially on where seeds are deposited in the environment. Some authors suggest that in forest understory seed predation is lower than in gaps, and higher than at the forest edge. However, most studies have been carried out in large forest patches and very little is known about the effects of microhabitat conditions on seed predation in forest fragments. We evaluated the effects of three microhabitats (gaps, forest edge, and understory) on seed predation of two palm species (Euterpe edulis and Syagrus romanzoffiana) in two semi-deciduous forest fragments (230 and 2100 ha) in southeast Brazil. Our objective was to test two hypotheses: (1) Low rodent abundance in small fragments as a result of meso-predator action levels leads to lower seed predation in small fragments. (2) Most mammal species in small fragments are generalists with respect to diet and habitat, so that seed predation is similar in different microhabitats (gaps, forest edge and understory) in the small fragment, but not in the larger one. The study community of small fragments is usually composed of generalist species (in diet and habitat aspects), so we expected the same rate of seed predation among microhabitats (gaps, forest edge and understory) in the tested smaller fragment. The experiment was carried out in the dry season (for E. edulis) and in the wet season (for S. romanzoffiana) in 1999. We conclude that post-dispersal seed predation in forest fragments can be directly connected with mammal communities, reflecting their historical and ecological aspects.     

Habitat use under latent predation risk. A case study with wintering forest birds

We test whether the spatial distribution of birds within a habitat is determined by predation risk and also by interspecific competition. The work was carried out in a montane mixed forest of central Spain with four Parus species, the long-tailed tit ( Aegithalos caudatus ) and the nuthatch ( Sitta europaea ). Experimental feeders, that varied in their risk of predation, were used to control the effect of natural variations in food availability and quality on the habitat use patterns of different species. Tree gleaning passerines avoided feeding on dark inner forest places far from edges, distant from protective cover, outside the inner tree canopy and near the ground; they preferred deciduous, relatively clear forest plots. These effects remained invariable across years and weather conditions. There was a common pattern of selection of foraging locations by the four Parus species: proximity to cover and height above ground and over the lowest branches of the tree canopy positively influenced the use of feeding places. According to these patterns, the vigilance proportion of species was significantly higher when feeding far from cover than when birds were feeding near pine foliage. This pattern was also common for the four studied Parus species. Nevertheless, the interspecific dominance hierarchy of the species was positively correlated with the use of the safest feeders (feeders farther the ground and nearer from protective cover within tree canopy), being the converse with the most exposed ones. Therefore, the results of this paper demonstrate that the selection of feeding locations within habitat follows a pattern minimizing predation risk. Interspecific dominance hierarchies can lead to the exploitation of unfavourable risky patches by subordinate species.     

Density-dependent mortality of wintering Dunlins Calidris alpina through predation by Eurasian Sparrowhawks Accipiter nisus

Density-dependent mortality in wintering shorebirds is critical to shorebird population dynamics, but is difficult to study and demonstrate. Most winter mortality of small shorebirds probably results from predation by raptors. Over 13 winters, I studied Eurasian Sparrowhawk Accipiter nisus predation on wintering Dunlins Calidris alpina at a rocky shore in south-east Scotland, a site apparently of marginal suitability for wintering Dunlins. Most Dunlins on the study site were juveniles (first winter birds), and winter juvenile mortality rate due to Sparrowhawk predation was strongly and positively related to Dunlin density (maximum winter count). There was no indication of any marked effect of weather on Dunlin winter mortality rate, except that mortality was slightly greater in winters with lower temperatures. Dunlin mortality was unrelated to the mortality through Sparrowhawk predation of a preferred prey, the Redshank Tringa totanus , or to sighting rates of hawks, suggesting that increased hunting of Redshanks by hawks or a numerical response by hawks were not involved in density-dependent mortality of Dunlins.     

Differential mortality of wintering shorebirds on the Banc d'Arguin, Mauritania, due to predation by large falcons

Predators may influence many aspects of the daily life and seasonal movements of their prey. Here we quantify direct, and evaluate indirect effects of predation by three falcon species (Lanner Falcon Falco biarmicus , Barbary Falcon Falco pelegrinoides and Peregrine Falcon Falco peregrinus ) on coastal shorebirds wintering on the Banc d'Arguin, Mauritania, an area hosting approximately 30% of the East Atlantic Flyway population of shorebirds. On the basis of 754 h of observation over five winters, 97 witnessed attacks and 585 collected prey remains, we show that shorebirds were safer in larger flocks, which tended to be attacked less often. Furthermore, species that forage relatively close to shore and in small flocks were depredated more often than expected from their relative abundance. In three species, Red Knot Calidris canutus canutus , Bar-tailed Godwit Limosa lapponica taymyrensis and Dunlin Calidris alpina , the juveniles were more vulnerable than adults. We estimated that on average 1% of the juvenile and 0.1% of the adult Red Knots present were killed by large falcons each winter. For Red Knots we simultaneously quantified annual survival on the basis of an individual colour-marking programme: mortality due to predation by falcons accounted for an estimated 6.2% (juveniles) and 0.8% (adults) of annual mortality. We suggest that juvenile Red Knots are 10 times as likely to be killed by falcons because they use riskier habitats, i.e. early and late tide foraging areas closer to shores where surprise attacks are both more common and more successful. These results indicate that the strength of indirect effects of predation operating in a shorebird population largely outweigh the effects of mortality per se .