Mo A. Verhoeven

When Siberia came to the Netherlands : The response of continental black-tailed godwits to a rare spring weather event

1. Extreme weather events have the potential to alter both short- and long-term population dynamics as well as community- and ecosystem-level function. Such events are rare and stochastic, making it difficult to fully document how organisms respond to them and predict the repercussions of similar events in the future. 2. To improve our understanding of the mechanisms by which short-term events can incur long-term consequences, we documented the behavioural responses and fitness consequences for a long-distance migratory bird, the continental black-tailed godwit Limosa limosa limosa, resulting from a spring snowstorm and three-week period of record low temperatures. 3. The event caused measurable responses at three spatial scales - continental, regional and local - including migratory delays (+19 days), reverse migrations (>90 km), elevated metabolic costs (+8·8% maintenance metabolic rate) and increased foraging rates (+37%). 4. There were few long-term fitness consequences, however, and subsequent breeding seasons instead witnessed high levels of reproductive success and little evidence of carry-over effects. 5. This suggests that populations with continued access to food, behavioural flexibility and time to dissipate the costs of the event can likely withstand the consequences of an extreme weather event. For populations constrained in one of these respects, though, extreme events may entail extreme ecological consequences.

Estimating the size of the Dutch breeding population of continental black-tailed godwits from 2007-2015 using resighting data from spring staging sites

Over the past 50 years, the population of Continental Black-tailed Godwits Limosa limosa limosa breeding of the East Atlantic Flyway has been in steep decline. This decline has previously been documented in trend analyses and six Netherlands-wide count-based population estimates, the last of which was completed in 1999. We provide an updated population size estimate and describe inter-annual fluctuations in the population between 2007 and 2015. To generate these estimates, we integrated a mark-recapture survival analysis with estimates of the densities of colour-marked individuals at migratory staging sites with known proportions of Continental and Icelandic L. l. islandica Black-tailed Godwits within a Bayesian framework. The use of these analytical techniques means that, in contrast with earlier efforts, our estimates are accompanied with confidence intervals, allowing us to estimate the population size with known precision. Using additional information on the breeding destination of 43 godwits equipped with satellite transmitters at Iberian staging areas, we found that 87% (75–95% 95% CI) of the nominate subspecies in the East Atlantic Flyway breed in The Netherlands. We estimated that the number of breeding pairs in The Netherlands has declined from 47,000 (38,000–56,000) pairs in 2007 to 33,000 (26,000–41,000) in 2015. Despite a temporary increase in 2010 and 2011, the population declined by an average of 3.7% per year over the entire period from 2007–2015, and by 6.3% from 2011–2015. We conclude that investing in an intensive demographic programme at a regional scale, when combined with targeted resightings of marked individuals elsewhere, can yield population estimates at the flyway scale.

Does wintering north or south of the Sahara correlate with timing and breeding performance in black-tailed godwits?

Abstract Migrating long distances requires time and energy, and may interact with an individuals performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.

Fuelling and moult in Red Knots before northward departure: a visual evaluation of differences between ages, sexes and subspecies

Abstract The departure of migratory birds from their non-breeding grounds is thought to be driven by the phenology of their breeding destination. In north-west Australia, two plumage morphs of Red Knot (Calidris canutus) prepare for a 5500-kmjourney to Yellow Sea staging areas. These morphs are recognised as the subspecies C. c. piersmai and C. c. rogersi, which breed at different latitudes and have different seasonalities. From February to May 2011, we observed the migratory preparation of individually marked birds of known age, sex and type. This enabled a comparison of fuelling rates and prealternate moult among these classes. First-year birds did not prepare for migration. Second-year birds accumulated smaller fuel stores and reached lower plumage scores than adults. Adults of both types reached their highest abdominal profile scores by the end of April when they were last observed in Roebuck Bay. This lack of difference between types in the timing of fuelling and departure is surprising. Based on the differences in staging and breeding phenology, C. c. rogersi is expected to leave north-west Australia 2–4 weeks before C. c. piersmai. Assuming that types and subspecies are equivalent, our findings in combination with other research on Red Knots in the East Asian-Australasian Flyway suggest that it takes more than breeding origin alone to explain annual cycles in migratory birds. Concurrent migratory schedules imply that, during northward staging in the Yellow Sea, there is strong variation in fuelling rates between and within subspecies depending on non-breeding origin. The ongoing loss of staging habitat may therefore have differential effects on Red Knots in the East Asian-Australasian Flyway.

High-altitude shorebird migration in the absence of topographical barriers: avoiding high air temperatures and searching for profitable winds

Nearly 20% of all bird species migrate between breeding and nonbreeding sites annually. Their migrations include storied feats of endurance and physiology, from non-stop trans-Pacific crossings to flights at the cruising altitudes of jetliners. Despite intense interest in these performances, there remains great uncertainty about which factors most directly influence bird behaviour during migratory flights. We used GPS trackers that measure an individuals altitude and wingbeat frequency to track the migration of black-tailed godwits (Limosa limosa) and identify the abiotic factors influencing their in-flight migratory behaviour. We found that godwits flew at altitudes above 5000 m during 21% of all migratory flights, and reached maximum flight altitudes of nearly 6000 m. The partial pressure of oxygen at these altitudes is less than 50% of that at sea level, yet these extremely high flights occurred in the absence of topographical barriers. Instead, they were associated with high air temperatures at lower altitudes and increasing wind support at higher altitudes. Our results therefore suggest that wind, temperature and topography all play a role in determining migratory behaviour, but that their relative importance is context dependent. Extremely high-altitude flights may thus not be especially rare, but they may only occur in very specific environmental contexts.

Generational shift in spring staging site use by a long-distance migratory bird

In response to environmental change, species have been observed altering their migratory behaviours. Few studies, however, have been able to determine whether these alterations resulted from inherited, plastic or flexible changes. Here, we present a unique observation of a rapid population-level shift in migratory routes—over 300 km from Spain to Portugal—by continental black-tailed godwits Limosa limosa limosa. This shift did not result from adult godwits changing staging sites, as adult site use was highly consistent. Rather, the shift resulted from young godwits predominantly using Portugal over Spain. We found no differences in reproductive success or survival among individuals using either staging site, indicating that the shift resulted from developmental plasticity rather than natural selection. Our results therefore suggest that new migratory routes can develop within a generation and that young individuals may be the agents of such rapid changes.

Warming springs and habitat alteration interact to impact timing of breeding and population dynamics in a migratory bird

In seasonal environments, increasing spring temperatures lead many taxa to advance the timing of reproduction. Species that do not may suffer lower fitness. We investigated why black-tailed godwits (Limosa limosa limosa), a ground-breeding agricultural grassland shorebird, have not advanced timing of reproduction during the last three decades in the face of climate change and human-induced habitat degradation. We used data from an 11-year field study to parameterize an Integral Projection Model to predict how spring temperature and habitat quality simultaneously influence the timing of reproduction and population dynamics. We found apparent selection for earlier laying, but not a correlation between the laying dates of parents and their offspring. Nevertheless, in warmer springs, laying dates of adults show a stronger positive correlation with laying date in previous springs than in cooler ones, and this leads us to predict a slight advance in the timing of reproduction if spring temperatures continue to increase. We also show that only in landscapes with low agricultural activity, the population can continue to act as a source. This study shows how climate change and declining habitat quality may enhance extinction risk.

Primary moult of continental Black-tailed Godwits Limosa limosa limosa in the Doñana wetlands, Spain

ABSTRACT Capsule: Most Continental Black-tailed Godwits Limosa limosa limosa using the Doñana wetlands during post-breeding migration appear to begin moult before they arrive and suspend moult before they migrate onwards to West Africa. Aims: We aim to describe the primary moult strategies and patterns in the Continental Black-tailed Godwits using the Doñana wetlands, a major passage and wintering area for waterbirds in southern Spain. Methods: Individual godwits were captured, marked and their primary moult was scored in Doñana during the non-breeding season (June–March) in 2011 and 2012. Data from resightings of colour-marked godwits and birds equipped with satellite transmitters were used to estimate stopover duration during post-breeding migration (June–September) to determine if godwits move to West Africa before completing their primary moult. Results: Average primary moult duration was estimated to be 84 days ± 9 se, during 29 June–21 September and did not differ between sexes. Only 2% of individuals were observed with suspended moult. We estimated stopover duration in Doñana to be 13 days ± 2 se before migrating to West Africa. Conclusions: Most godwits stage for about two weeks in the Doñana wetlands during southward migration, moult their primaries and appear to suspend moult before crossing the Sahara. Others may complete their primary moult in Doñana, or elsewhere in Europe and overwinter in Doñana where increasing numbers of godwits have been detected in recent years. A few individuals may finish the moult in Doñana and migrate to West Africa late in the post-breeding season.