Philip W. Atkinson

Rapid population decline in red knots: fitness consequences of decreased refuelling rates and late arrival in Delaware Bay

Most populations of migrant shorebirds around the world are in serious decline, suggesting that vital condition–dependent rates such as fecundity and annual survival are being affected globally. A striking example is the red knot (Calidris canutus rufa) population wintering in Tierra del Fuego, which undertakes marathon 30 000 km hemispheric migrations annually. In spring, migrant birds forage voraciously on horseshoe crab eggs in Delaware Bay in the eastern USA before departing to breed in Arctic polar deserts. 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 from concurrent over–harvesting of crabs. Reduced nutrient storage, especially in late–arriving birds, possibly combined with reduced sizes of intestine and liver during refuelling, had severe fitness consequences for adult survival and recruitment of young in 2000–2002. From 1997 to 2002 known survivors in Delaware Bay were heavier at initial capture than birds never seen again, annual survival of adults decreased by 37% between May 2000 and May 2001, and the number of second–year birds in wintering flocks declined by 47%. Population size in Tierra del Fuego declined alarmingly from 51 000 to 27 000 in 2000–2002, seriously threatening the viability of this subspecies. Demographic modelling predicts imminent endangerment and an increased risk of extinction of the subspecies without urgent risk–averse management.

The buffer effect and large-scale population regulation in migratory birds.

Buffer effects occur when sites vary in quality and fluctuations in population size are mirrored by large changes in animal numbers in poor-quality sites but only small changes in good-quality sites. Hence, the poor sites ‘buffer’ the good sites, a mechanism that can potentially drive population regulation if there are demographic costs of inhabiting poor sites. Here we show that for a migratory bird this process can apply on a country-wide scale with consequences for both survival and timing of arrival on the breeding grounds (an indicator of reproductive success). The Icelandic population of the black-tailed godwit, Limosa limosa islandica, wintering in Britain has increased fourfold since the 1970s (ref. 5) but rates of change within individual estuaries have varied from zero to sixfold increases. In accordance with the buffer effect, rates of increase are greater on estuaries with low initial numbers, and godwits on these sites have lower prey-intake rates, lower survival rates and arrive later in Iceland than godwits on sites with stable populations. The buffer effect can therefore be a major process influencing large-scale population regulation of migratory species.

Changes in commercially fished shellfish stocks and shorebird populations in the Wash, England

Abstract The Wash, in eastern England, supports internationally important populations of 11 species of shorebird. A major commercial shellfishery provides potential conflict between fishermen and nature conservation interests. During the 1990s, high fishing mortality and low recruitment substantially reduced the stocks of cockle (Cerastoderma edule) and mussel (Mytilus edulis). Population models, constructed from estimates of survival and recruitment, indicated that declines in the availability of cockles and mussels were associated with changes in oystercatcher (Haemotopus ostralegus) survival between 1970 and 1998, including three periods of mass mortality, and also the recruitment of juvenile birds to both oystercatcher and knot (Calidris canutus) populations. Emigration of knot may also have taken place. Cockle recruitment was dependent on climatic conditions, whereas mussel populations tended to be stable. The decline in mussel stocks, due to over-fishing, increased the vulnerability of the oystercatcher population to mass-mortality episodes in poor cockle years. The key to preventing major oystercatcher kills in future is to ensure sufficient mussels are available in poor cockle years. Recent cultivation of mussels in inter-tidal areas has been beneficial and is an important management tool for maintaining bird populations.

Conserving the Birds of Uganda’s Banana-Coffee Arc: Land Sparing and Land Sharing Compared

Reconciling the aims of feeding an ever more demanding human population and conserving biodiversity is a difficult challenge. Here, we explore potential solutions by assessing whether land sparing (farming for high yield, potentially enabling the protection of non-farmland habitat), land sharing (lower yielding farming with more biodiversity within farmland) or a mixed strategy would result in better bird conservation outcomes for a specified level of agricultural production. We surveyed forest and farmland study areas in southern Uganda, measuring the population density of 256 bird species and agricultural yield: food energy and gross income. Parametric non-linear functions relating density to yield were fitted. Species were identified as “winners” (total population size always at least as great with agriculture present as without it) or “losers” (total population sometimes or always reduced with agriculture present) for a range of targets for total agricultural production. For each target we determined whether each species would be predicted to have a higher total population with land sparing, land sharing or with any intermediate level of sparing at an intermediate yield. We found that most species were expected to have their highest total populations with land sparing, particularly loser species and species with small global range sizes. Hence, more species would benefit from high-yield farming if used as part of a strategy to reduce forest loss than from low-yield farming and land sharing, as has been found in Ghana and India in a previous study. We caution against advocacy for high-yield farming alone as a means to deliver land sparing if it is done without strong protection for natural habitats, other ecosystem services and social welfare. Instead, we suggest that conservationists explore how conservation and agricultural policies can be better integrated to deliver land sparing by, for example, combining land-use planning and agronomic support for small farmers.

Declines in British-breeding populations of Afro-Palaearctic migrant birds are linked to bioclimatic wintering zone in Africa, possibly via constraints on arrival time advancement

Capsule The bioclimatic zone and habitat type within which birds winter are the most important determinants of population trends. Aims To investigate whether regional factors on wintering grounds, phenological mismatch, or habitat on breeding or wintering grounds show relationships with population changes of Afro-Palaearctic migrant birds. Methods We modelled breeding bird survey trends of 26 species of Afro-Palaearctic migrant birds that breed in Britain, and assessed the most important variables. We also investigated spatial variation in population trends within the UK. Results Bioclimatic zone and wintering habitat type were the most important determinants of population trend. After accounting for the effects of the other variables, species that spend the winter in southern Africa or the humid tropics of central and western Africa showed more negative population trends than those that winter in the arid zone. Species occupying open and woodland habitats declined and generalist species increased, on average. We also suggest that the effect of bioclimatic zone on population trends may be through constraints on the timing of migration. Conclusion Correlations between population change and both wintering area and winter habitat suggest regional changes in climate or land-use in the humid tropics are driving declines in many long-distance migrant species, possibly partly through migratory constraints.

Demographic consequences of migratory stopover: linking red knot survival to horseshoe crab spawning abundance

Understanding how events during one period of the annual cycle carry over to affect survival and other fitness components in other periods is essential to understanding migratory bird demography and conservation needs. Previous research has suggested that western Atlantic red knot (Calidris canutus rufa) populations are greatly affected by horseshoe crab (Limulus polyphemus) egg availability at Delaware Bay stopover sites during their spring northward migration. We present a mass-based multistate, capture-recapture/resighting model linking (1) red knot stopover mass gain to horseshoe crab spawning abundance and (2) subsequent apparent annual survival to mass state at the time of departure from the Delaware Bay stopover area. The model and analysis use capture-recapture/resighting data with over 16,000 individual captures and 13,000 resightings collected in Delaware Bay over a 12 year period from 1997–2008, and the results are used to evaluate the central management hypothesis that red knot populations can be influenced by horseshoe crab harvest regulations as part of a larger adaptive management effort. Model selection statistics showed support for a positive relationship between horseshoe crab spawning abundance during the stopover and the probability of red knots gaining mass (parameter coefficient from the top model b = 1.71, = 0.46). Our analyses also supported the link between red knot mass and apparent annual survival, although average estimates for the two mass classes differed only slightly. The addition of arctic snow depth as a covariate influencing apparent survival improved the fit of the data to the models (parameter coefficient from the top model b = 0.50, = 0.08). Our results indicate that managing horseshoe crab resources in the Delaware Bay has the potential to improve red knot population status.

Numbers of wintering waterbirds in Great Britain and the Isle of Man (1994/1995–1998/1999): II. Coastal waders (Charadrii)

Abstract Great Britains wintering coastal wader populations have been estimated for the period 1994/1995–1998/1999 from data provided from two sources: the Wetland Bird Survey (WeBS) and the Non-estuarine Coastal Waterfowl Survey (UK-NEWS). New methodology for estimating the number of wintering waders is applied. It imputes (fills in) the value of missing counts before estimating the population size of each species as being the mean of the largest annual count made between November and March over the relevant 5-year period. This methodology has led to an 11% larger estimate of the number of waders present on Great Britains coasts than the traditional approach based on averaging just January counts over a 5-year period, and it suggests that Great Britains coastline supports ca. 2.1 million waders. Updated values are presented from which site evaluations based on 1% of the national population can be derived. Great Britain is of considerable international importance for waders. It holds >25 and >50% of the flyway populations of nine and four species of wader, respectively, but for the first time since the start of monitoring in the early 1970s, the historical increase in the number of predominantly coastal waders wintering in Great Britain is coming to an end. Seven of the 14 species that have shown population changes of >5% since the last set of 1987/1988–1991/1992 population estimates have declined in numbers. The possible causes of the fluctuations in wader populations, such as climate change and changing nutrient inputs to coastal waters are discussed. There is an urgent need to identify the causative factors leading to these declines, and to use the new population estimates to identify new sites that should be afforded legal protection, an action that should help Great Britain maintain its internationally important wader populations. The decrease in the updated population estimate of Eurasian oystercatcher, for example, has made it possible to determine that 19 rather than 17 sites in Great Britain are worthy of statutory protection on the basis of holding 1% or more of its national population.

Population decline is linked to migration route in the Common Cuckoo

Migratory species are in rapid decline globally. Although most mortality in long-distance migrant birds is thought to occur during migration, evidence of conditions on migration affecting breeding population sizes has been completely lacking. We addressed this by tracking 42 male Common Cuckoos from the rapidly declining UK population during 56 autumn migrations in 2011–14. Uniquely, the birds use two distinct routes to reach the same wintering grounds, allowing assessment of survival during migration independently of origin and destination. Mortality up to completion of the Sahara crossing (the major ecological barrier encountered in both routes) is higher for birds using the shorter route. The proportion of birds using this route strongly correlates with population decline across nine local breeding populations. Knowledge of variability in migratory behaviour and performance linked to robust population change data may therefore be necessary to understand population declines of migratory species and efficiently target conservation resources.

Rapid and continued population decline in the Spoon-billed Sandpiper Eurynorhynchus pygmeus indicates imminent extinction unless conservation action is taken

Summary The Spoon-billed Sandpiper Eurynorhynchus pygmeus (IUCN Category: Critically Endangered) is in rapid decline. Data from across the entire breeding range (Chukotka and Koryakya in the Russian far north-east) and especially from the well-studied southern core breeding area at Meinypilgyno, confirm the continuing strong decline. At four breeding sites, where more than two counts were available for analysis, the decline was estimated at 26% per annum between 2002 and 2009 ,o r an 88% decline over this period. Allowing for unsurveyed areas, this equates to a decline from a total population of approximately 1,000 breeding pairs in 2000 to 120–220 in 2009. Breeding studies at Meinypilgyno in 2003–2007 (not 2006) showed that the proportion of nests hatching at least one chick was 0.65 and once chicks left the nest, the mean brood size of chicks up to one week old was 1.99. Where it was possible to follow broods, 0.61 chicks fledged per nesting attempt. Survival and recruitment analysis of birds ringed at Meinypilgyno indicated that annual adult survival did not significantly differ over the 2003–2009 study but that recruitment in to the adult breeding population was effectively zero in all but one year of the study (2005). Resighting data for the last two years of the study were sparse due to very low numbers of marked adults being recorded and survival rates over the last 2–3 years of the study must therefore be treated with caution. The analysis therefore indicated that after fledging, survival during immaturity must be very low, leading to a low (or no) recruitment into an ageing population. Recent observations collated from the non-breeding areas confirm the declining trend observed in the breeding areas and imply that the main threats to the population lie along the migration route or in the wintering areas. These are poorly known although hunting in the wintering areas has been identified as a major mortality factor. Other threats include major loss of their intertidal habitats, and collection of birds on the breeding areas by specimen collectors. Improved monitoring in both the breeding and non-breeding areas as well as research on juvenile survival is recommended. Concerted international conservation action is essential if this species is to avoid extinction. This requires (i) improved understanding of the main wintering and staging areas and associated threats; (ii) addressing those threats that can be tackled with immediate effect, such as hunting; (iii) continued long-term monitoring on the breeding areas; (iv) an exploration of other potential breeding areas; (v) conservation action at all important stop-over and wintering sites along the entire flyway and (vi) consideration of a captive-breeding programme to ensure the survival of this species.

Staging Behavior in Red KNOT (Calidris Canutus) in Delaware BAY: Implications for Monitoring Mass and Population Size

Abstract.— Many migratory birds use staging sites to gain essential resources to fuel their ongoing migration. Understanding staging strategies reveals much about migration systems and is essential if one is concerned with monitoring population trends and mass gains, two of the principal methods for assessing the “health” of a migratory population. In spring 2004, we investigated the staging behavior in Delaware Bay of Red Knot (Calidris canutus) using mark-recapture techniques and resightings of birds marked in the preceding spring. Individuals staged for 11–12 days, which declined to 8–10 days late in the season. Arrivals were asynchronous, but departures tended to be synchronized. A simple sensitivity analysis showed that the mark-recapture analysis estimated length of stay to within +10% and confirmed biases in monitoring trends and mass gains using peak counts and mass-on-date regressions. Alternative methods using staging duration to estimate passage population size and mass gains were shown to be unbiased. Using these methods, we estimated a passage population size in 2004 of 18,000 Red Knot that arrived at an average mass of 111 g and, on average, gained mass at 7.2 g day-1. Thus, in 2004, the passage population was substantially smaller than the recent peak count of 50,360 in 1998, which confirms a significant decline in the number of Red Knot staging in Delaware Bay. Use of refined techniques such as these is essential if management decisions such as those in Delaware Bay are to be based on firm scientific advice.