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Migratory shorebirds are among the most threatened groups of birds. They rely on natural intertidal habitats outside the breeding season, but, to some extent have adjusted to using man-made habitats. Here, we assessed the importance of coastal saltpans – a type of anthropogenic wetland – for feeding in migratory shorebirds during their northward migration along the East Asian-Australasian Flyway (EAAF). We combined low tide counts on intertidal flats and nearby saltpans at the Luannan coastal wetland complex (Bohai Bay, China) with Bayesian mixing model analyses (BMMs) based on stable isotopes to evaluate the relative importance of coastal saltpans versus natural intertidal habitats as foraging grounds for migrating species. We grouped shorebird species (n = 24) according to feeding guild and body size, and found that both predictors explained the broad-scale patterns of foraging use of saltpans by shorebirds at low tide. The guild of water-surface foraging species (e.g. stilts and avocets), independently of body size, mostly fed in saltpans, and the small-medium visual (e.g. plovers) and tactile-surface (e.g. sandpipers) foraging species consumed a significant portion of their diet in this habitat. In contrast, most large tactile-surface foraging species barely foraged in saltpans at low tide. BMMs showed that shorebirds had a greater reliance on saltpans than did traditional counts of foraging birds in each habitat at low tide. Saltpan food is rich in essential fatty acids, so the contribution of saltpans to the diet of shorebirds should not be considered only in absolute values, but also in the quality of this contribution. Saltpans may therefore help conserve declining shorebirds if properly managed – for example by controlling water levels – to serve the specific feeding guilds that rely on them. While our focus is in the EAAF, the findings are relevant for other flyways and other non-tidal anthropogenic wetlands.

The Bohai Black-tailed Godwit (Limosa limosa bohaii) is a newly discovered subspecies in the East Asian-Australasian Flyway. Based on satellite tracking of 21 individuals that were tagged in northern Bohai Bay, China, from 2016 to 2018, we here describe the annual cycle of this subspecies. All the birds had Thailand as their southernmost ‘winter’ destination. The spring departure was in late March during northward migration, Bohai Bay was the first stopping site where they spent on average 39 days (± SD = 6 d), followed by Inner Mongolia and Jilin province (stopping for 8 d ± 1 d). The arrival of the breeding grounds in the Russian Far East was centred in late May. Two breeding sites were detected, with average locations 1100 km apart; the eastern site was beyond the known Asian breeding distribution of the Black-tailed Godwit. Southward migration started in late June, with the godwits tending to make longer stops at the same two main stopping sites used in the spring, i.e. Inner Mongolia and Jilin province (32 ± 5 d) and Bohai Bay (44 ± 8 d), with some individuals making a third stop in the middle-lower reaches of the Yangtze River in southern China (12 ± 4 d). By the end of September, most tracked individuals had arrived in Thailand. Compared with the previously known subspecies, bohaii godwits have strikingly different schedules of migration and moult, this study thus adding to the knowledge about intraspecific diversity of black-tailed godwits in the East Asian-Australian Flyway.

Populations can rapidly respond to environmental change via adaptive phenotypic plasticity, which can also modify interactions between individuals and their environment, affecting population dynamics. Bird migration is a highly plastic resource-tracking tactic in seasonal environments. However, the link between the population dynamics of migratory birds and migration tactic plasticity is not well-understood. The quality of staging habitats affects individuals' migration timing and energy budgets in the course of migration and can consequently affect individuals' breeding and overwintering performance, and impact population dynamics. Given staging habitats being lost in many parts of the world, our goal is to investigate responses of individual migration tactics and population dynamics in the face of loss of staging habitat and to identify the key processes connecting them. We started by constructing and analysing a general full-annual-cycle individual-based model with a stylized migratory population to generate hypotheses on how changes in the size of staging habitat might drive changes in individual stopover duration and population dynamics. Next, through the interrogation of survey data, we tested these hypotheses by analysing population trends and stopover duration of migratory waterbirds experiencing the loss of staging habitat. Our modelling exercise led to us posing the following hypotheses: the loss of staging habitat generates plasticity in migration tactics, with individuals remaining on the staging habitat for longer to obtain food due to a reduction in per capita food availability. The subsequent increasing population density on the staging habitat has knock-on effects on population dynamics in the breeding and overwintering stage. Our empirical results were consistent with the modelling predictions. Our results demonstrate how environmental change that impacts one energetically costly life-history stage in migratory birds can have population dynamic impacts across the entire annual cycle via phenotypic plasticity.种群因具备适应性的表型可塑性可以迅速响应环境变化，同时，表型可塑性也能够通过改变个体与其环境之间的相互作用而影响种群动态。迁徙是鸟类在季节性变化的环境中演化出的一种资源追踪型策略，其可塑性非常强。然而有关迁徙策略可塑性与候鸟种群动态之间的反馈过程尚不清楚。 迁徙中停地的质量影响着候鸟个体在迁徙过程中时间与能量的权衡，进而通过影响个体在繁殖和越冬阶段的表现来影响种群动态。由于世界范围内很多迁徙鸟类的中停地都在急剧减少，因此本文希望探究在迁徙中停地丧失的情况下，鸟类的迁徙策略与种群动态之间存在何种联结。 首先，我们构建了一个包含程式化迁徙种群全年周期的基于个体的模型，利用该模型提出了关于迁徙中停地的变化如何驱动候鸟个体停留时间和种群动态发生改变的假设，并通过对中国渤海湾地区迁徙中停的水鸟的调查数据的分析进行了验证。 我们的假设是：迁徙中停地的丧失会导致鸟类迁徙策略的改变，由于候鸟个体在中停地可获得的食物资源减少，它们会延长在中停地的停留时间以补充能量，由此引发中停地鸟类种群密度的增长，从而对整个生活史周期内候鸟繁殖和越冬阶段的种群动态产生了连锁反应。我们的实证研究结果与建模预测一致。 本研究表明，环境变化对迁徙候鸟生活史中某一个阶段的作用会通过表型可塑性对其整个周期的种群动态产生巨大影响。.

Occurring across Eurasia, the Black-tailed GodwitLimosa limosahas three recognized subspecies,melanuroides,limosaandislandicafrom east to west, respectively. With the smallest body size,melanuroideshas been considered the only subspecies in the East Asian-Australasian Flyway. Yet, observations along the Chinese coast indicated the presence of distinctively large individuals. Here we compared the morphometrics of these larger birds captured in northern Bohai Bay, China, with those of the three known subspecies and explore the genetic population structuring of Black-tailed Godwits based on the control region of the mitochondrial genome (mtDNA). We found that the Bohai Godwits were indeed significantly larger thanmelanuroides, resemblinglimosamore thanislandica, but with relatively longer bills thanislandica. The level of genetic differentiation between Bohai Godwits and the three recognized subspecies was of similar magnitude to the differentiation among previously recognized subspecies. Based on these segregating morphological and genetic characteristics, we propose that these birds belong to a distinct population, which may be treated and described as a new subspecies.

Sex ratio is a fundamental concept in evolutional biology, and theory predicts that parents should invest in sons and daughters according to the fitness returns they expect from them. The fitness returns may depend on the timing of breeding and on parental conditions leading to sex ratios that depend on breeding date and/or parental quality. Here, we investigate the offspring sex ratio in a small shorebird, the Kentish Plover Charadrius alexandrinus, in a large breeding population in Eastern China, and test whether the parents adjust their offspring’s sex in response to hatch date, brood age and their own body condition. Using 1264 chicks from 676 broods that were molecularly sexed, we show that hatchling sex ratio was not significantly different from unity. Hatchling sex ratios were not related to hatch date or to the body condition of parents. In addition, we sexed 138 eggs that were confiscated from illegal egg collectors and found that the mortality of female and male embryos was not significantly different. The latter result is important by suggesting that neither primary sex ratio (i.e., at conception) nor secondary sex ratio (i.e., at hatching) is biased. Taken together, the even offspring sex ratio in Chinese Kentish Plovers is consistent with recent analyses of six plover populations that found even sex ratios at hatching. Future works should investigate whether the even sex ratio persists into adulthood, or it may shift toward more males (or females) due to sex-biased mortalities of juveniles and/or adults.

SummaryThe 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 km2) 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 km2) at high and low tide and on the adjacent natural tidal flats (57 km2) 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.

Background Coastal saltpans are a common supratidal human-modified wetland habitat found within many coastal landscape mosaics. Commercial salt production and aquaculture practices often result in the creation of exposed coastal substrates that could provide suitable breeding habitat for waterbird populations; however, few studies have quantified waterbird breeding success in these artificial wetlands. Methods Here we examine the nesting behavior of the Gull-billed tern (Gelochelidon nilotica) breeding in the Nanpu coastal saltpans of Bohai Bay, Yellow Sea, China over three consecutive nesting seasons (2017–2019) by using nest survival model in Program MARK. Results The results revealed that nest survival of Gull-billed terns in coastal saltpans (0.697) was higher than previously published estimates from other regions, with an estimated daily survival rate (DSR) of 0.982 ± 0.001 (±95% CI). High nest survival was mainly attributed to low levels of human disturbances and low predation rates, while exposure to strong winds, flooding and silting were the main factors causing nest failure. Model-averaged estimates revealed that eggs laid in nests located on ‘habitat islands’ with feather or clam shell substrates were most likely to hatch. Initiation date, nest age, clutch size and quadratic effects of nearest-neighbor distance, nearest distance to road and nearest distance to water were all significant predictors of nest success, but the nest survival declined overall from 2017 to 2019 due to the degradation and loss of breeding habitat anthropogenically caused by rising water levels. Discussion Coastal saltpans represent an alternative breeding habitat for the Gull-billed tern populations in Bohai Bay, but conservation management should prioritize flood prevention to improve the extent and quality of breeding habitat, concurrent with efforts to create further ‘habitat islands’ with suitable nesting substrate.

Loss and degradation of wetlands has occurred worldwide, impacting ecosystems and contributing to the decline of waterbirds, including shorebirds that occur along the heavily developed coasts of the East Asian-Australasian Flyway (EAAF). Artificial (i.e. human-made) wetlands are pervasive in the EAAF and known to be used by shorebirds, but this phenomenon has not been systematically reviewed. We collated data and expert knowledge to understand the extent and intensity of shorebird use of coastal artificial habitats along the EAAF. We found records of 83 species, including all regularly occurring coastal migratory shorebirds, across 176 artificial sites with eight different land uses. Thirty-six species including eleven threatened species occurred in internationally important numbers. However, threatened species were less likely to occur, and larger-bodied, migratory and coastal specialist species less likely to feed, at artificial sites. Abundance, species richness and density varied across artificial habitats, with high abundance and richness but low density on salt production sites; high abundance and density on port and power production sites; and, low abundance and richness on aquaculture and agriculture. Overall, use of coastal artificial habitats by shorebirds is widespread in the flyway, warranting a concerted effort to integrate artificial habitats alongside natural wetlands into conservation frameworks. Salt production sites are cause for particular concern because they support large shorebird aggregations but are often at risk of production cessation and conversion to other land uses. Preserving and improving the condition of all remaining natural habitats and managing artificial habitats are priorities for shorebird conservation in the EAAF [see Supplementary Materials A for a Japanese translation of the abstract].

Understanding and resolving conflicts between phenotypic and genetic differentiation is central to evolutionary research. While phenotypically monomorphic species may exhibit deep genetic divergences, some morphologically distinct taxa lack notable genetic differentiation. Here we conduct a molecular investigation of an enigmatic shorebird with a convoluted taxonomic history, the White-faced Plover (Charadrius alexandrinus dealbatus), widely regarded as a subspecies of the Kentish Plover (C. alexandrinus). Described as distinct in 1863, its name was consistently misapplied in subsequent decades until taxonomic clarification ensued in 2008. Using a recently proposed test of species delimitation, we reconfirm the phenotypic distinctness of dealbatus. We then compare three mitochondrial and seven nuclear DNA markers among 278 samples of dealbatus and alexandrinus from across their breeding range and four other closely related plovers. We fail to find any population genetic differentiation between dealbatus and alexandrinus, whereas the other species are deeply diverged at the study loci. Kentish Plovers join a small but growing list of species for which low levels of genetic differentiation are accompanied by the presence of strong phenotypic divergence, suggesting that diagnostic phenotypic characters may be encoded by few genes that are difficult to detect. Alternatively, gene expression differences may be crucial in producing different phenotypes whereas neutral differentiation may be lagging behind.