Tamar Lok

Simultaneous declines in summer survival of three shorebird species signals a flyway at risk

Summary 1. There is increasing concern about the world’s animal migrations. With many land-use and climatological changes occurring simultaneously, pinning down the causes of large-scale conservation problems requires sophisticated and data-intensive approaches. 2. Declining shorebird numbers along the East Asian–Australasian Flyway, in combination with data on habitat loss along the Yellow Sea (where these birds refuel during long-distance migrations), indicate a flyway under threat. 3. If habitat loss at staging areas indeed leads to flyway-wide bird losses, we would predict that: (i) decreases in survival only occur during the season that birds use the Yellow Sea, and (ii) decreases in survival occur in migrants that share a reliance on the vanishing intertidal flats along the Yellow Sea, even if ecologically distinct and using different breeding grounds. 4. Monitored from 2006–2013, we analysed seasonal apparent survival patterns of three shorebird species with non-overlapping Arctic breeding areas and considerable differences in foraging ecology, but a shared use of both north-west Australian non-breeding grounds and the Yellow Sea coasts to refuel during northward and southward migrations (red knot Calidris canutus piersmai, great knot Calidris tenuirostris, bar-tailed godwit Limosa lapponica menzbieri). Distinguishing two three-month non-breeding periods and a six-month migration and breeding period, and analysing survival of the three species and the three seasons in a single model, we statistically evaluated differences at both the species and season levels. 5. Whereas apparent survival remained high in north-west Australia, during the time away from the non-breeding grounds survival in all three species began to decline in 2011, having lost 20 percentage points by 2012. By 2012 annual apparent survival had become as low as 0� 71 in bar-tailed godwits, 0� 68 in great knots and 0� 67 in red knots. In a separate analysis for red knots, no mortality occurred during the migration from Australia to China. In the summers of low summer survival, weather conditions were benign in the Arctic breeding areas. 6. We argue that rapid seashore habitat loss in the Yellow Sea is the most likely explanation of reduced summer survival, with dire (but uncertain) forecasts for the future of these flyway populations. This interpretation is consistent with recent findings of declining shorebird numbers at seemingly intact southern non-breeding sites.

Body shrinkage due to Arctic warming reduces red knot fitness in tropical wintering range

Consequences conferred at a distance Migratory animals have adapted to life in multiple, sometimes very different environments. Thus, they may show particularly complex responses as climates rapidly change. Van Gils et al. show that body size in red knot birds has been decreasing as their Arctic breeding ground warms (see the Perspective by Wikelski and Tertitski). However, the real toll of this change appears not in the rapidly changing northern part of their range but in the apparently more stable tropical wintering range. The resulting smaller, short-billed birds have difficulty reaching their major food source, deeply buried mollusks, which decreases the survival of birds born during particularly warm years. Science, this issue p. 819; see also p. 775 A warming Arctic decreases the fitness of migratory red knots in their distant wintering range. Reductions in body size are increasingly being identified as a response to climate warming. Here we present evidence for a case of such body shrinkage, potentially due to malnutrition in early life. We show that an avian long-distance migrant (red knot, Calidris canutus canutus), which is experiencing globally unrivaled warming rates at its high-Arctic breeding grounds, produces smaller offspring with shorter bills during summers with early snowmelt. This has consequences half a world away at their tropical wintering grounds, where shorter-billed individuals have reduced survival rates. This is associated with these molluscivores eating fewer deeply buried bivalve prey and more shallowly buried seagrass rhizomes. We suggest that seasonal migrants can experience reduced fitness at one end of their range as a result of a changing climate at the other end.

The cost of migration: spoonbills suffer higher mortality during trans-Saharan spring migrations only

Explanations for the wide variety of seasonal migration patterns of animals all carry the assumption that migration is costly and that this cost increases with migration distance. Although in some studies, the relationships between migration distance and breeding success or annual survival are established, none has investigated whether mortality during the actual migration increases with migration distance. Here, we compared seasonal survival between Eurasian spoonbills (Platalea leucorodia leucorodia) that breed in The Netherlands and migrate different distances (ca 1000, 2000 and 4500 km) to winter in France, Iberia and Mauritania, respectively. On the basis of resightings of individually marked birds throughout the year between 2005 and 2012, we show that summer, autumn and winter survival were very high and independent of migration distance, whereas mortality during spring migration was much higher (18%) for the birds that wintered in Mauritania, compared with those flying only as far as France (5%) or Iberia (6%). As such, this study is the first to show empirical evidence for increased mortality during some long migrations, likely driven by the presence of a physical barrier (the Sahara desert) in combination with suboptimal fuelling and unfavourable weather conditions en route.

Declining adult survival of New Zealand Bar-tailed Godwits during 2005–2012 despite apparent population stability

Abstract Like many migratory shorebird populations using the East Asian—Australasian Flyway, Bar-tailed Godwits Limosa lapponica baueri in New Zealand have significantly declined since the mid-1990s, but census data indicate a relatively stable population since 2004. The demographic drivers of both the decline and stabilisation remain unknown. We estimated annual survival from mark—recapture data of adult godwits in New Zealand during 2005–2014. Annual adult survival declined over the study period from 0.89-0.96 in 2005–2010 to 0.83-0.84 in 20112012. The simultaneous decline in annual survival found in a separate study of Bar-tailed Godwits L. l. menzbieri in north-west Australia suggests a common effect of their high dependence on threatened migratory staging sites in the Yellow Sea; the more extreme decline in L. l. menzbieri may reflect ecological differences between the populations, such as timing and extent of use of these sites. At current apparent recruitment rates, persistent adult survival of ∼0.84 would lead to a population decline of 5–6% per year in L. l. baueri. Our study implies that the demographic precursors to a population decline developed during a period of apparent population stability; this suggests that monitoring a single index of population stability is insufficient for predicting future trends.

Seasonal variation in density dependence in age‐specific survival of a long‐distance migrant

Density dependence in vital rates is key to population regulation. Rather than being constant, the strength of density dependence may vary throughout the year, but empirical evidence is limited. Based on 22 years of data of color-banded birds from a recovering population of Eurasian Spoonbills Platalea leucorodia leucorodia, we show, for the first time, seasonal variation in density dependence in survival of a long-distance migrating bird. Combining resightings and dead recoveries at breeding, stopover, and nonbreeding areas enabled us to (1) separate true survival from permanent emigration from the breeding area, and (2) estimate survival in three seasons: summer, early winter (including autumn migration), and late winter (including spring migration). Accompanying the rapid population growth, juvenile annual survival initially increased, manifested in early winter, but thereafter, at high population sizes, it strongly decreased through a combination of decreasing survival in all seasons. Annual survival of subadult (second- and third-year) and adult birds decreased more gradually with increasing population size, with density dependence occurring in early winter for subadults and late winter for adults. Thus, the shape and strength of density dependence in survival varied with age and season. Understanding the seasonal timing of density dependence, especially with reference to underlying mechanisms, is important for the design of effective conservation strategies.

Migration Tendency Delays Distributional Response to Differential Survival Prospects along a Flyway

When populations grow or decline, habitat selection may change due to local density-dependent processes, such as site dependence and interference. In seasonally migrating animals, nonbreeding distributions may be determined through these mechanisms of density dependence, which we examine here at a hemispheric scale for a long-distance migrating bird. Using summer and winter resightings of 2,095 Eurasian spoonbills Platalea leucorodia leucorodia that were ringed in the Netherlands during 16 years of fast population growth, we show that neither site dependence nor interference fully explains their patterns of survival and winter distribution. Within their three main wintering areas, annual survival decreased with an increase in population size. While survival was consistently higher in the two European wintering areas (France, Iberia), most spoonbills migrated onward to winter in west Africa. The number of birds wintering in Europe increased, but not enough to maximize annual survival. We conclude that a constraint of tradition (their “migration tendency”) inhibits birds from changing their migratory habits. We pose that this phenomenon may similarly constrain other migratory populations from rapidly responding to large-scale climate- and/or human-driven habitat changes at their wintering grounds.

Interpreting variation in growth of Eurasian spoonbill chicks : Disentangling the effects of age, sex and environment

Chick body condition can be a sensitive indicator of local environmental conditions and has been shown to be correlated with chick survival. Designing a reliable index of chick body condition for a given species from a single measurement point requires knowledge about the extent of variation in body size, about chick age and about the relative sensitivity of the growth of different biometric measures to variation in environmental conditions. To gain this knowledge, we describe sex-specific variation in growth of several morphometric measures and body mass of Eurasian Spoonbills Platalea leucorodia. We repeatedly measured 35 chicks that grew up in small colonies on the island of Schiermonnikoog to derive detailed growth curves until fledging (based on the 12 surviving chicks) and to assess the extent of reduction in growth of starved chicks measured at least twice (n = 11) compared to those that survived. Growth curves until fledging were compared with biometric measurements of two to five week old chicks from (mostly) larger colonies of which hatching date was accurately estimated (n = 631). Growth of all measures, except the eighth primary, was sex-specific, with the most pronounced sex effect on the asymptotic values of tarsus length and body mass: adult males were predicted to become 17% heavier than females and to have 22% longer tarsi than females. Body mass and tarsus growth tended to be more reduced under food deprivation than (head-)bill and eighth primary growth. As an index of chick body condition, we propose to use the proportional deviation in body mass from the predicted body mass for a given age and sex. To do so, measurements of nearly fledged Spoonbill chicks should include at least eighth primary length to estimate age, tarsus length to estimate sex, and body mass as a measure that integrates age, sex and environmental effects.