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Annual routines of migratory birds inferred from archival solar geolocation devices have never before been confirmed using GPS technologies. A female black-tailed godwit Limosa limosa limosa captured on the breeding grounds in the Netherlands in 2013 and recaptured in 2014 was outfitted with both an Intigeo geolocator and an UvA-BiTS GPS-tracker. The GPS positions show that, after its breeding season in 2013, the godwit flew 2035 km nonstop from the Netherlands to southern Spain. It then spent the entire nonbreeding season in the southern part of the Iberian Peninsula before returning to the Netherlands the following spring, stopping for 7 days in the delta of the Ebro River in Spain, and again for a day in central Belgium. To compare the geolocation and GPS data, we analysed the geolocation data with two open-source software packages: one using a threshold method (GeoLight) and the other a template-fit approach (FLightR). Estimates using GeoLight, on average, deviated from the individual’s true position by 495.5 ± 1031.2 km (great circle distance with equinoxes excluded), while FLightR estimates deviated by 43.3 ± 51.5 km (great circle distance with equinoxes included). Arrival and departure schedules estimated by FLightR were within 12 h of those determined by the GPS tracker, whereas GeoLight’s estimates were less precise. For the analysed track, FLightR represents an improvement over GeoLight; if true for other species and conditions, FLightR will hopefully help establish more precise and accurate uses of geolocation data in tracking studies. To aid future improvements in the analysis of solar geolocation data, we also provide the GPS and geolocation data files together with our R scripts as Supplementary material Appendix 1–6.

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. 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. 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. 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. 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. 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. Policy implications. Due to established economic interests, governments are usually reluctant to act for conservation, unless unambiguous evidence for particular cause–effect chains is apparent. This study adds to an increasing body of evidence that habitat loss along the Yellow Sea shores explains the widespread declines in shorebird numbers along the East Asian–Australasian Flyway and threatens the long‐term prospects of several long‐distance migrating species. To halt further losses, the clearance of coastal intertidal habitat must stop now.

1. Intensification of agricultural use of grassland habitats has been linked to the declines of many farmland bird species, several of whom have been in decline for multiple decades despite agri-environmental schemes. In the Netherlands, where most grasslands have been transformed into well-drained monocultures managed for maximal dairy production, schemes that aim to protect nests from agricultural activities are the most popular. The cause of the failure of these schemes, however, seems poor recruitment of farmland birds. 2. Using the black-tailed godwit Limosa limosa limosa as our model species, we tested the hypothesis underlying the most popular agri-environmental schemes that there are no differences in condition at hatch, growth, and apparent survival rates between chicks raised on monocultures compared with chicks from herb-rich meadows with high water tables, often managed, for the sake of grassland birds, in traditional ways. We also compare the growth rates from both habitat types with those of chicks from dairy farmland from 1976 to 1985, a time when population declines of godwits first became visible. 3. Hatchling mass did not differ between chicks from herb-rich meadows and grassland monocultures, but chicks hatched on monocultures were on average 14—16% lighter at fledging, and had 4% smaller bills than chicks hatched on herb-rich meadows. The growth rates of female chicks hatched on herb-rich meadows were similar to those of chicks measured from 1976 to 1985. Males fledged lighter and smaller than females. 4. Apparent survival during the first year of life for chicks hatched on herb-rich meadows was 2·5 times higher than that of chicks hatched on grassland monocultures. The apparent survival on herb-rich meadows seems sufficient for a stable population. 5. Synthesis and applications: We found striking differences between chicks hatched on modern grassland monocultures and herb-rich meadows. That chicks hatched on monocultures had lower growth and survival rates than chicks on meadows indicates that these chicks suffer a higher risk of starvation and/or predation. These findings imply that the most often applied agri-environmental schemes (i.e. payments per clutch found and postponed mowing), are not effective. Instead, these schemes could even encourage maladaptive habitat choice. Conservation efforts should thus focus on the provision of herb-rich meadows with high groundwater tables.

Background The degradation and loss of critical nodes in the East Asian–Australasian Flyway (EAAF) migration network could have implications for the migration of shorebirds. The conservation of critical nodes along the EAAF is important for maintaining the stability of migratory networks. However, identifying these nodes remains a major challenge. Methods We used a network method that integrates GPS tracking data and remote sensing data to identify critical nodes for three shorebird species (Black-tailed Godwit ( Limosa limosa ), Eurasian Curlew ( Numenius arquata ), and Pied Avocet ( Recurvirostra avosetta )) in migratory networks along the EAAF and priority conservation areas to facilitate the development of conservation and management plans. Results A total of 111, 37, and 81 nodes in the migratory networks were identified for the Black-tailed Godwit, Eurasian Curlew, and Pied Avocet, respectively; these included 25, 13, and 16 critical nodes, respectively. Node importance (node betweenness) was higher in the migration period than in the breeding and wintering periods. Habitat loss was observed in most stopover nodes. A total of 23 unprotected critical nodes were identified as priority conservation areas. Conclusions Habitat loss was observed in most of the unprotected critical nodes in stopover regions for all three species. The inclusion of these sites such as Haoyao sumu forest farm and Tianjin coast, into flyway site networks (the East Asian–Australasian Flyway Partnership (EAAFP)) should be prioritized, and the habitat quality of nearby alternative nodes should be improved. Future studies should focus on developing new node indexes that integrate ecological variables with each node’s role in maintaining network connectivity using data from a larger number of tracked individuals and birdwatching data. Such studies could help identify currently unknown regions with critical habitats.

Whether to disperse, and where to, are two of the most prominent decisions in an individual's life, with major consequences for reproductive success. We studied natal and breeding dispersal in the monogamous black‐tailed godwit Limosa limosa limosa in the Netherlands, where they breed in agricultural grasslands. The majority of these grasslands recently changed from wet herb‐rich meadows into well‐drained grassland monocultures, on which godwits have a lower reproductive success. Here we examine habitat selection with a multistate mark–recapture analysis. Habitat transition probabilities between meadows and monocultures were estimated on the basis of 1810 marked chicks and 531 adults during seven years in a 8500 ha study area. Young and adult godwits may differ in habitat selection because: 1) adults may have gained experience from previous nest success where to settle, 2) younger individuals may find it harder to compete for the best territories. Both young and adults moved at a higher rate from the predominant monocultures to meadows than the other way around, thus actively selecting the habitat with better quality. However, dispersal distance of adults was not affected by previous nest success. The average dispersal distance from place of birth of godwits breeding for the first time was ten times larger than that of adult godwits. That godwits breeding in their second calendar year arrived and laid at similar dates and were equally able to select territories in areas with high breeding densities, suggests that young birds were not competitively inferior to adults. Although on monocultures reproduction is insufficient to maintain constant populations, birds sometimes moved from meadows to monocultures. This explains why even after 30 years of land‐use intensification, godwits still breed in low‐quality habitat. The adjustment to changing habitat conditions at the population level appears to be a slow process.

Migrating long distances requires time and energy, and may interact with an individual's 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. In this manuscript, we show that individuals of a long‐distance migrating shorebird (the black‐tailed godwit Limosa limosa) which spend their nonbreeding period in West Africa arrived and laid earlier than godwits migrating to wintering locations north of the Sahara, mainly in Doñana Wetlands, Spain. This contrasts with previously studied migrating bird species where migration distance tends to be positively correlated with arrival and/or laying date. In spite of their late arrival and laying, godwits which wintered closer to their Dutch breeding grounds laid larger eggs. We could, however, not find a correlation with other reproductive parameters.

On the basis of morphological differences, three subspecies of Black-tailed Godwit (Limosa limosa) have been recognized (L. l. limosa, L. l. islandica and L. l. melanuroides). In previous studies mitochondrial DNA (mtDNA) sequence data showed minimal genetic divergence between the three subspecies and an absence of sub-structuring within L. l. limosa. Here, population genetic structure and phylogeographic patterns have been analyzed using COI, HVR1 and HVR2 mtDNA sequence data as well as 12 microsatellite loci (nuDNA). The nuDNA data suggest genetic differentiation between L. l. limosa from Sweden and The Netherlands, between L. l. limosa and L. l. islandica, but not between L. l. limosa and L. l. melanuroides. However, the mtDNA data were not consistent with the nuDNA pattern. mtDNA did support a split between L. l. melanuroides and L. l. limosa/L. l. islandica and also demonstrated two L. l. limosa haplotype clusters that were not geographically isolated. This genetic structure can be explained by a scenario of isolation of L. l. melanuroides from L. l. limosa in Beringia during the Last Glacial Maximum. During the Pleistocene separation of L. l. islandica from L. l. limosa occurred, followed by colonization of Iceland by the L. l. islandica during the Holocene. Within L. l. limosa founder events, followed by population expansion, took place during the Holocene also. According to the patterns observed in both markers together and their geographic separation, we propose that the three traditional subspecies indeed represent three separate genetic units.

The natural coastal wetlands of the East Asian-Australasian Flyway (EAAF) are disappearing at alarming rates, leading to rapid declines of many populations of waterbirds in the most species-rich flyway in the world. The identification and assessment of possible alternative habitats that may buffer the loss of natural wetlands should, therefore, be a priority for the conservation of migratory waterbirds using this flyway. Coastal saltpans are functional wetlands that support large numbers of waterbirds worldwide. The Nanpu Saltpans in the northern Bohai Bay of the Yellow Sea in China are one of the largest (290 km 2 ) saltpan complexes in the world. In this paper, we document the value of the Nanpu Saltpans for supporting waterbirds. The surveys, carried out from 2013 to 2016, included waterbird counts in the saltpans (93 km 2 ) at high and low tide and on the adjacent natural tidal flats (57 km 2 ) at low tide. Of the 89 waterbird species recorded, 27 had maximum counts exceeding the 1% threshold value of estimated flyway populations. The maximum counts of waterbirds in northward migration and southward migration in the Nanpu Saltpans were 96,000 and 93,500, respectively, including both foraging and roosting birds; these figures do not account for turnover, so the total number of birds using the site is likely to be higher. The maximum counts on the adjacent tidal flats at low tide amounted to 73,000 and 20,000 waterbirds during northward and southward migration, respectively, and most of them were foraging birds. In the boreal winter, few birds fed in the saltpans, but several thousand fed on the tidal flats. Waterbirds used the inland ponds (2.0–18.0 km from the intertidal area) mainly for feeding both during low tide and high tide and used the nearshore ponds (0.3–4.3 km from the intertidal area) mainly for high-tide roosting. Some species, such as Black-tailed Godwit Limosa limosa, Marsh Sandpiper Tringa stagnatilis, Pied Avocet Recurvirostra avosetta, and Black-winged Stilt Himantopus himantopus, occurred mainly in the saltpans; other species preferred tidal flats, such as Red Knot Calidris canutus, Great Knot Calidris tenuirostris, Bar-tailed Godwit Limosa lapponica, Eurasian Curlew Numenius arquata, Relict Gull Larus relictus, and Grey Plover Pluvialis squatarola. This study clearly demonstrates the joint ecological function of the Nanpu Saltpan complex and adjacent tidal flats as a key staging area for waterbirds in the EAAF, and as such both urgently warrant protected status.

Shorebirds (Charadriiformes) undergo rapid migrations with potential for long‐distance dispersal (LDD) of plants. We studied the frequency of endozoochory by shorebirds in different parts of Europe covering a broad latitudinal range and different seasons. We assessed whether plants dispersed conformed to morphological dispersal syndromes. A total of 409 excreta samples (271 faeces and 138 pellets) were collected from redshank Tringa totanus, black‐winged stilt Himantopus himantopus, pied avocet Recurvirostra avosetta, northern lapwing Vanellus vanellus, Eurasian curlew Numenius arquata and black‐tailed godwit Limosa limosa in south‐west Spain, north‐west England, southern Ireland and Iceland in 2005 and 2016, and intact seeds were extracted and identified. Godwits were sampled just before or after migratory movements between England and Iceland. The germinability of seeds was tested. Intact diaspores were recovered from all bird species and study areas, and were present in 13% of samples overall. Thirteen plant families were represented, including Charophyceae and 26 angiosperm taxa. Only four species had an ‘endozoochory syndrome’. Four alien species were recorded. Ellenberg values classified three species as aquatic and 20 as terrestrial. Overall, 89% of seeds were from terrestrial plants, and 11% from aquatic plants. Average seed length was higher in redshank pellets than in their faeces. Six species were germinated, none of which had an endozoochory syndrome. Seeds were recorded during spring and autumn migration. Plant species recorded have broad latitudinal ranges consistent with LDD via shorebirds. Crucially, morphological syndromes do not adequately predict LDD potential, and more empirical work is required to identify which plants are dispersed by shorebirds. Incorporating endozoochory by shorebirds and other migratory waterbirds into plant distribution models would allow us to better understand the natural processes that facilitated colonization of oceanic islands, or to improve predictions of how plants will respond to climate change, or how alien species spread.

Migrating birds make the longest non‐stop endurance flights in the animal kingdom. Satellite technology is now providing direct evidence on the lengths and durations of these flights and associated staging episodes for individual birds. Using this technology, we compared the migration performance of two subspecies of bar‐tailed godwit Limosa lapponica travelling between non‐breeding grounds in New Zealand (subspecies baueri) and northwest Australia (subspecies menzbieri) and breeding grounds in Alaska and eastern Russia, respectively. Individuals of both subspecies made long, usually non‐stop, flights from non‐breeding grounds to coastal staging grounds in the Yellow Sea region of East Asia (average 10 060 ± SD 290 km for baueri and 5860 ± 240 km for menzbieri). After an average stay of 41.2 ± 4.8 d, baueri flew over the North Pacific Ocean before heading northeast to the Alaskan breeding grounds (6770 ± 800 km). Menzbieri staged for 38.4 ± 2.5 d, and flew over land and sea northeast to high arctic Russia (4170 ± 370 km). The post‐breeding journey for baueri involved several weeks of staging in southwest Alaska followed by non‐stop flights across the Pacific Ocean to New Zealand (11 690 km in a complete track) or stopovers on islands in the southwestern Pacific en route to New Zealand and eastern Australia. By contrast, menzbieri returned to Australia via stopovers in the New Siberian Islands, Russia, and back at the Yellow Sea; birds travelled on average 4510 ± 360 km from Russia to the Yellow Sea, staged there for 40.8 ± 5.6 d, and then flew another 5680–7180 km to Australia (10 820 ± 300 km in total). Overall, the entire migration of the single baueri godwit with a fully completed return track totalled 29 280 km and involved 20 d of major migratory flight over a round‐trip journey of 174 d. The entire migrations of menzbieri averaged 21 940 ± 570 km, including 14 d of major migratory flights out of 154 d total. Godwits of both populations exhibit extreme flight performance, and baueri makes the longest (southbound) and second‐longest (northbound) non‐stop migratory flights documented for any bird. Both subspecies essentially make single stops when moving between non‐breeding and breeding sites in opposite hemispheres. This reinforces the critical importance of the intertidal habitats used by fuelling godwits in Australasia, the Yellow Sea, and Alaska.