Differences in food abundance cause inter-annual variation in the breeding phenology of High Arctic waders

Previous work has shown that High Arctic waders in Greenland are “income breeders”, i.e. the resources used for egg formation are based almost entirely on biomass obtained on the breeding grounds. Thus, their breeding phenology is expected to be highly sensitive to inter-annual variation in food abundance during the pre-laying period. Early spring snow-cover may also influence timing of egg-laying either directly or mediated through food resources. Here, we report on the inter-annual variation in clutch initiation of three wader species breeding in High Arctic Greenland, Sanderling (Calidris alba), Dunlin (Calidris alpina) and Ruddy Turnstone (Arenaria interpres), in relation to spring snow-cover and spring arthropod abundance over ten breeding seasons at Zackenberg Research Station 1995–2005. Food abundance had the strongest effect on timing of clutch initiation, while the proportion of snow-free land had a weaker but still significant effect, i.e. more food and more snow-free land both result in earlier egg-laying. We hypothesize that food is most important when there is sufficient snow-free land to nest on, while snow-cover is of increasing importance in years with late snowmelt.     

Diurnal variability in the detection of waders on their breeding grounds

When censusing breeding waders in large upland areas, the time of day has a major influence on the numbers of birds detected.     

Testing the effects of recreational disturbance on two upland breeding waders

Previous work has shown that Golden Plovers Pluvialis apricaria guarding chicks utilize heavily disturbed habitat at a lower rate than surrounding areas, but that such rates of avoidance are reduced when the movement of people is restricted to a surfaced footpath. However, there remained some unanswered questions, which are addressed in this paper. First, we examine to what extent habitat avoidance is dependent upon visitor numbers, and find no evidence that Golden Plovers avoided disturbed areas at a site where visitor pressure was half that previously studied. Secondly, we examine whether these relationships between habitat occupancy and disturbance apply to other upland waders and find that Dunlin Calidris alpina habitat utilization in disturbed areas showed a non-significant increase of approximately 50% following the provision of a surfaced footpath, in a manner similar to that observed for Golden Plover. Thirdly, we examine whether the large numbers of visitors (120 per weekend day) using the surfaced footpath impact on Golden Plover breeding success, despite the lack of habitat avoidance. There was no evidence that nest location, clutch survival or chick growth rates were reduced close to the footpath. Together, these results suggest that high levels of disturbance can impact upon habitat usage by upland waders, but only in limited circumstances where visitor pressure is very high (greater than at least 30 visitors per weekend day). However, access to such areas can be permitted for large numbers of visitors without impacting upon wader reproductive performance through the provision of a well-surfaced route.     

The Swedish Arctic charr breeding programme

Results and experiences from a selective breeding programme aiming at improving the performance of an Arctic charr strain in aquaculture are presented. The programme, which has been running since 1985, uses traditional quantitative genetic methods based on relatedness and trait measurements. Traits considered included growth, age at sexual maturity, flesh colour, fat content and other features. Estimates in the early phase of the programme showed promising heritabilities for most traits, e.g. growth heritabilities of 0.34–0.52, and suggested that improvement by selection was feasible. Several traits such as a high rate of early maturation and poor flesh colour that were considered problematic by the Arctic charr farming industry at the programme’s start no longer hamper farming. Considerable improvement of growth by selection, estimated at 8% per generation, has contributed to shortening the production cycle in commercial farming. Results from studies of genotype–environment interactions are presented and discussed. Poor survival of fertilised eggs is a major problem in Swedish Arctic charr farming as indicated by presented survival rates. Efforts have consistently been made to avoid inbreeding in the selected strain, and at the present seventh generation the accumulated increase in inbreeding is estimated from pedigree data to be only approximately 5%.     

Arctic waders and the capital‐income continuum: Further tests using isotopic contrasts of egg components

Birds migrating annually to high‐latitude breeding grounds may benefit from the transport of endogenous nutrient reserves that ultimately contribute to reproduction. Shorebirds represent a diverse group of Arctic breeders that typically arrive on the breeding grounds with body reserves enriched in ¹³C and ¹⁵N due to wintering and staging in marine or estuarine habitats. Such isotopic differences between endogenous macronutrient reserves and local foodwebs allow the use of stable isotopes to test for the source of nutrient allocations to eggs. We examined δ¹³C and δ¹⁵N values in lipid‐free yolk and albumen and δ¹³C values in yolk lipid of first clutches of ten species of sandpiper and plover breeding near Churchill, Manitoba, Canada in 2003. Most birds had egg isotope values indicating a C3 terrestrial biome, which fits primarily an income (exogenous) breeding strategy. Two exceptions were single sandpiper and plover with strong marine isotope values. Among species, strong positive relationships for each isotope between egg tissue components suggest that egg proteins and lipids tended to be derived from the same isotopic source. Correlations of egg δ¹³C values for lipids and proteins approached theoretical relationships expected for exogenous breeding strategies, based on captive studies. Significant positive correlations between clutch initiation date and δ¹³C values of egg lipids and albumen suggest some endogenous nutrient contributions to later laid eggs but the circumstances under which this may occur are unstudied. Where possible, we recommend that researchers use blood and fat biopsies from laying females as a means of anchoring endogenous and exogenous endpoints for modeling of each reproductive event. We encourage the isotopic analysis of egg albumen, yolk and yolk lipids among individuals and species and tests of correlations among these components as a means of inferring origins of nutrients to eggs.     

Incubation scheduling by Northern Fulmars ( Fulmarus glacialis ) in the Canadian High Arctic

Northern Fulmars (Fulmarus glacialis) are a ubiquitous seabird of the North Atlantic and North Pacific oceans that breed in colonies surrounded by markedly different marine environmental conditions. I have studied the incubation behavior of fulmars at a remote colony in the Canadian High Arctic, where the birds had to cross 200 km of sea-ice during the early incubation period to reach feeding areas. These fulmars completed incubation in fewer shifts and had a longer mean incubation shift duration (5.3 days) than their counterparts breeding in the Boreal oceanographic zone. In particular, the mean duration of the first incubation shift by males in successful pairs (10.3 days) was 2 days longer than that for males in unsuccessful pairs and was longer than that reported at any other colony. This exceptionally long shift by the male may be required at this site to give females enough time to recoup energetic reserves after egg-laying, at a time when marine productivity in the Arctic is still seasonably low.     

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.     

The effects of improvement of upland, marginal grasslands on the breeding success of Lapwings Vanellus vanellus and other waders

Data on the breeding success of Lapwings Vanellus vanellus on unimproved and improved grassland, with comparative data for arable land, were obtained from a study on 760 Lapwing clutches. The study was undertaken in the Eden Valley, Cumbria, and Teesdale, Co. Durham, between 1985 and 1987. First clutches were larger on unimproved grassland with a mean of 3.73 eggs compared to 3.61 eggs on improved grassland. In contrast, replacement clutches were larger on improved areas (3.90 eggs) than on unimproved (3.47 eggs). On average, 40% of eggs laid on unimproved pastures hatched compared to only 17% on improved pastures. No significant difference in hatching success was found between unimproved and improved meadows with 32% and 22% of eggs hatching, respectively. Overall, 73% of unsuccessful first clutches were replaced on unimproved pastures, whereas on meadows and improved pastures combined, only 32% were replaced. Survival of small chicks was highest on unimproved areas. Production on unimproved areas was sufficient to replace adult losses and so maintain numbers, whereas on improved land production was too low to maintain existing breeding densities. Hatching success on unimproved areas was similar for the four species of wader considered. On improved areas, Redshank hatching success was relatively high, with 54% of pairs producing chicks, compared to 35% for Lapwing and 23% for Curlew.     

The use of predator-exclusion fencing as a management tool improves the breeding success of waders on lowland wet grassland

Waders breeding on lowland wet grassland have undergone dramatic declines across Europe in recent decades. Few species now achieve the levels of breeding success required for population stability and recovery, with predation from large mammals acting as a key compounding factor limiting nest survival and productivity. Predator management through lethal control is often controversial, yet alternative non-lethal methods are little tested in the context of grassland breeding waders. Excluding predators through the use of electric fences has led to improvements in nest and chick survival in other habitats. To test the applicability of this method to lowland wet grassland we constructed predator-exclusion fences on sites across the UK and, with Lapwing Vanellus vanellus as a study species, used historical and contemporary data to test whether excluding large mammalian predators leads to an increase in wader nest survival and productivity, and whether effects differ between fence designs. Lapwing nest survival was significantly higher in the presence of any type of predator-exclusion fence, with significantly fewer nests predated each day. Overall productivity also improved, with significantly higher numbers of chicks fledged per pair in years when fences were operational. Different designs and methods of powering fences resulted in different levels of success, with combination design fences and those powered by mains electricity performing best. Excluding large mammalian predators from areas of lowland wet grassland with predator-exclusion fencing successfully improves Lapwing nest survival and productivity, allowing breeding success to exceed the levels required for population recovery. Other wader species breeding in the same habitat are also likely to benefit from the increased protection from predation provided by fences. Predator-exclusion fencing is therefore an effective management tool for protecting restricted and declining populations of breeding waders on lowland wet grassland.     

Diet of black guillemots and northern fulmars breeding beside a High Arctic polynya

In the Arctic, the availability of early-season open water (shore leads, polynyas) is a key factor influencing annual reproduction by marine birds, as these relatively productive locations provide the only sites where migrating and breeding birds can feed. We examined the diet of two marine birds, black guillemots (Cepphus grylle) and northern fulmars (Fulmarus glacialis), feeding in the Hell Gate–Cardigan Strait Polynya in the Canadian High Arctic during the breeding seasons of 1980–1984. There was little overlap in prey items consumed between these two species, except that both relied on similar sympagic amphipods. Guillemots fed principally on benthic prey, and compared to earlier High Arctic studies, they consumed proportionally more benthic fish, mysids and decapods. Fulmars appeared to feed on swarms of pelagic or surface prey, and in this polynya they consumed proportionally fewer fish and more polychaetes than has been reported for fulmars at other High Arctic locations. Given that climate change may alter marine food webs by changing the types and availability of prey, our results set a baseline of dietary information for these predators against which future conditions can be compared.     

Phenological variation in annual timing of hibernation and breeding in nearby populations of Arctic ground squirrels

Ecologists need an empirical understanding of physiological and behavioural adjustments that animals can make in response to seasonal and long-term variations in environmental conditions. Because many species experience trade-offs between timing and duration of one seasonal event versus another and because interacting species may also shift phenologies at different rates, it is possible that, in aggregate, phenological shifts could result in mismatches that disrupt ecological communities. We investigated the timing of seasonal events over 14 years in two Arctic ground squirrel populations living 20 km apart in Northern Alaska. At Atigun River, snow melt occurred 27 days earlier and snow cover began 17 days later than at Toolik Lake. This spatial differential was reflected in significant variation in the timing of most seasonal events in ground squirrels living at the two sites. Although reproductive males ended seasonal torpor on the same date at both sites, Atigun males emerged from hibernation 9 days earlier and entered hibernation 5 days later than Toolik males. Atigun females emerged and bred 13 days earlier and entered hibernation 9 days earlier than those at Toolik. We propose that this variation in phenology over a small spatial scale is likely generated by plasticity of physiological mechanisms that may also provide individuals the ability to respond to variation in environmental conditions over time.     

Raptor predation on wintering waders in southeast Scotland

Predation on waders was investigated over the winters of 1982-83 and 1983-84 at a rocky shore and a small estuary in southeast Scotland. Probably all the waders which were found after being eaten were killed by raptors; mammalian predation and death due to severe weather were of minimal importance in comparison to the effects of raptor predation. Sparrowhawks were probably the major predators of waders at both sites, but owls also took some. Differences were found between the rocky shore and the estuary in the species of raptors which were seen (more Peregrines and Merlins seen on the estuary) and in the numbers of wader kills which had been plucked and eaten on open ground (more open ground pluckings, indicative of Peregrines and Merlins, found on the estuary). It is suggested that Peregrines and Merlins took more waders at the estuary due to their preference for hunting over large open expanses. Winter mortality rates due to predation varied in different species of waders: at the rocky shore the Redshank suffered the heaviest winter mortality (20% and 16%) whilst on the estuary the Ringed Plover had the highest winter mortality (19%). At both sites no large waders such as Curlews and Oystercatchers died as a result of predators and for these species other factors, such as severe weather or starvation, appeared to be a greater source of mortality. Significantly more juvenile Redshanks, Turnstones and, probably, Dunlin and Ringed Plovers were found eaten than was expected from the age-composition of their respective populations. In the Redshank and Turnstone several differences in the behaviour of juveniles and adults were discovered which may have made juveniles more vulnerable to predation.     

Eggs in the freezer: energetic consequences of nest site and nest design in Arctic breeding shorebirds

Birds construct nests for several reasons. For species that breed in the Arctic, the insulative properties of nests are very important. Incubation is costly there and due to an increasing surface to volume ratio, more so in smaller species. Small species are therefore more likely to place their nests in thermally favourable microhabitats and/or to invest more in nest insulation than large species. To test this hypothesis, we examined characteristics of nests of six Arctic breeding shorebird species. All species chose thermally favourable nesting sites in a higher proportion than expected on the basis of habitat availability. Site choice did not differ between species. Depth to frozen ground, measured near the nests, decreased in the course of the season at similar non-species-specific speeds, but this depth increased with species size. Nest cup depth and nest scrape depth (nest cup without the lining) were unrelated to body mass (we applied an exponent of 0.73, to account for metabolic activity of the differently sized species). Cup depth divided by diameter(2) was used as a measure of nest cup shape. Small species had narrow and deep nests, while large species had wide shallow nests. The thickness of nest lining varied between 0.1 cm and 7.6 cm, and decreased significantly with body mass. We reconstruct the combined effect of different nest properties on the egg cooling coefficient using previously published quantitative relationships. The predicted effect of nest cup depth and lining depth on heat loss to the frozen ground did not correlate with body mass, but the sheltering effect of nest cup diameter against wind and the effects of lining material on the cooling coefficient increased with body mass. Our results suggest that small arctic shorebirds invest more in the insulation of their nests than large species.     

Intra-seasonal variation in zooplankton availability,chick diet and breeding performance of a high Arctic planktivorous seabird

Different phenological responses to climate changes by species representing preys and predators may lead to mismatch between functionally dependent components of an ecosystem, with important effects on its structure and functioning. Here, we investigate within-season variation in zooplankton availability, chick diet composition and breeding performance of a small planktivorous seabird, the little auk (Alle alle) in two large colonies in Hornsund and Magdalenefjorden, Spitsbergen, differing in synchrony of breeding (11-day vs. 22-day hatching period, respectively). Assuming similar zooplankton phenology and existing differences in duration of the little auk breeding period, we expected lower availability of the preferred food in the less synchronized colony in Magdalenefjorden and in consequence a negative effect on nestling body mass and survival. We found that in both colonies Calanus glacialis (copepodite stage CV) was the most important prey item in the chick diet making up 68–87 % of the biomass and energy of all prey items. The only exception was the end of the chick-rearing period in Magdalenefjorden, when contribution of this prey item was significantly lower (24–26 %). Thus, late breeders in Magdalenefjorden were apparently mismatched regarding C. glacialis CV availability. However, the hatching date did not affect birds fitness (reproductive output and chick pre-fledging mass) significantly. Results of our study indicate that little auks breeding on Spitsbergen can respond to a wide range of environmental conditions and prey availabilities through the plasticity of their foraging behaviour, which may help them to maintain their optimum fitness level in changing and unpredictable environments.