The onset of spring and timing of migration in two arctic nesting goose populations: the pink‐footed goose Anser bachyrhynchus and the barnacle goose Branta leucopsis

An earlier onset of spring has been recorded for many parts of Eurasia in recent decades. This has consequences for migratory species, both in changing the conditions encountered by individuals on reaching migratory sites and in affecting cues regulating the timing of migration where decisions to migrate are influenced by local environmental variables. Here we examine the timing of spring migration for two arctic goose populations, the pink‐footed goose Anser brachyrhynchus (during 1990–2003) and barnacle goose Branta leucopsis (during 1982–2003), which both breed on Svalbard. The satellite‐derived Normalised Difference Vegetation Index (NDVI) was used to express the onset of spring at their wintering and spring staging sites. Pink‐footed geese use several sites during spring migration, ranging from the southernmost wintering areas in Belgium to two spring staging areas in Norway, and distances between sites used along the flyway are relatively short. There was a positive correlation in the onset of spring between neighbouring sites, and the geese migrated earlier in early springs. Barnacle geese, on the other hand, have a long overseas crossing from their wintering grounds in Britain to spring staging areas in Norway. Although spring advanced in both regions, there was no corresponding correlation in the timing of onset of spring between their wintering and spring staging sites, and little evidence for barnacle geese migrating earlier over the whole study period. Hence, where geese can use spring conditions at one site as an indicator of the conditions they might encounter at the next, they have responded quickly to the advancement of spring, whereas in a situation where they cannot predict, they have not yet responded, despite the advancement of spring in the spring staging area.     

The consequences of climate-driven stop-over sites changes on migration schedules and fitness of Arctic geese

1. How climatic changes affect migratory birds remains difficult to predict because birds use multiple sites in a highly interdependent manner. A better understanding of how conditions along the flyway affect migration and ultimately fitness is of paramount interest. 2. Therefore, we developed a stochastic dynamic model to generate spatially and temporally explicit predictions of stop-over site use. For each site, we varied energy expenditure, onset of spring, intake rate and day-to-day stochasticity independently. We parameterized the model for the migration of pink-footed goose Anser brachyrhynchus from its wintering grounds in Western Europe to its breeding grounds on Arctic Svalbard. 3. Model results suggested that the birds follow a risk-averse strategy by avoiding sites with comparatively high energy expenditure or stochasticity levels in favour of sites with highly predictable food supply and low expenditure. Furthermore, the onset of spring on the stop-over sites had the most pronounced effect on staging times while intake rates had surprisingly little effect. 4. Subsequently, using empirical data, we tested whether observed changes in the onset of spring along the flyway explain the observed changes in migration schedules of pink-footed geese from 1990 to 2004. Model predictions generally agreed well with empirically observed migration patterns, with geese leaving the wintering grounds earlier while considerably extending their staging times in Norway.     

Spring migration patterns,habitat use,and stopover site protection status for two declining waterfowl species wintering in China as revealed by satellite tracking

East Asian migratory waterfowl have greatly declined since the 1950s, especially the populations that winter in China. Conservation is severely hampered by the lack of primary information about migration patterns and stopover sites. This study utilizes satellite tracking techniques and advanced spatial analyses to investigate spring migration of the greater white‐fronted goose (Anser albifrons) and tundra bean goose (Anser serrirostris) wintering along the Yangtze River Floodplain. Based on 24 tracks obtained from 21 individuals during the spring of 2015 and 2016, we found that the Northeast China Plain is far‐out the most intensively used stopover site during migration, with geese staying for over 1 month. This region has also been intensely developed for agriculture, suggesting a causal link to the decline in East Asian waterfowl wintering in China. The protection of waterbodies used as roosting area, especially those surrounded by intensive foraging land, is critical for waterfowl survival. Over 90% of the core area used during spring migration is not protected. We suggest that future ground surveys should target these areas to confirm their relevance for migratory waterfowl at the population level, and core roosting area at critical spring‐staging sites should be integrated in the network of protected areas along the flyway. Moreover, the potential bird–human conflict in core stopover area needs to be further studied. Our study illustrates how satellite tracking combined with spatial analyses can provide crucial insights necessary to improve the conservation of declining Migratory species.     

Expensive Traditions: Energy Expenditure of Aleutian Geese in Traditional and Recently Colonized Habitats

ABSTRACT In an effort to reduce goose depredation at a traditional spring migratory stopover site, private landowners implemented a coordinated hazing plan to scare Aleutian cackling geese (Branta hutchinsii leucopareia) from private lands to adjacent public pastures that were cultivated and set aside specifically for geese. Coincidentally, some Aleutian geese began using a new stopover site 150 km farther south in their spring migratory range; numbers at the new site continue to increase. We tested the idea that when their ability to acquire resources deteriorates geese are likely to seek improved foraging conditions, especially during spring migration when individuals strive to maximize nutrient stores and minimize energy expenditure. We quantified measures of goose foraging performance in traditional and new spring staging sites by calculating foraging opportunity, foraging effort, body condition, and daily energy expenditure. Geese staging at the site with higher levels of human disturbance had less foraging opportunity and, despite increased foraging effort and more nutritious food-plants at the site, birds there experienced an elevated energy expenditure and poorer body condition than birds at the new stopover site. Reduced foraging time and increased energy expenditure at the traditional spring staging site may have triggered the colonization process. Suitability assessment of habitat for migratory geese should include measures of foraging opportunity, disturbance risks, and daily energy expenditure in addition to quantity and quality of foods.     

Patch choice of avian herbivores along a migration trajectory–From Temperate to Arctic

Migratory waterfowl species seem to track temporal and spatial pulses of optimal forage availability on their way from temperate wintering to arctic breeding sites. In order to unravel the relative contribution of forage quality and forage biomass to foraging choices in avian herbivores, we experimentally manipulated biomass and quality of main forage plants through fertilisation and grazing exclusion at three sites along the flyway of barnacle geese, Branta leucopsis. Fertilisation increased the nitrogen content of the forage and grazing exclusion increased biomass levels. Manipulated plots were offered to wild geese in a random block experimental design and goose visitation was measured through dropping counts. At all sites there was a trend towards a higher preference of plots with increased quality and average biomass above plots with an average quality and increased biomass. Generally, geese preferred plots with highest standing crop of nitrogen. The numerical response of the geese to forage changes was supported by behavioural observations at the Baltic site. We conclude that for migrating barnacle geese the bottlenecks in the standing crop of nitrogen appear to lie in the limited biomass availability at the Baltic stopover site and the limited nutrient content of food in the Arctic breeding site, restricting the potential nutrient intake on these sites.     

Possible effects of local enrichment by gulls on feeding-site selection by wintering Barnacle Geese Branta leucopsis

The selection of salt-marsh feeding sites by wintering Barnacle Geese Branta leucopsis was investigated on Schiermonnikoog, The Netherlands. Use of sites by geese was estimated by counts of goose droppings. Geese spent significantly more time grazing in sites with evidence of Herring Gull Larus argentatus and Lesser Black-backed Gull Lfuscus breeding activity than in nearby non-gull sites. The nitrogen content of red fescue grass Festuca rubra was significantly greater in gull sites. We suggest that the geese preferred to feed in these locations because of the improved nutritional value of the grass. The higher nitrogen content of F. rubra in gull sites was most likely the result of nutrient inputs from gull droppings rather than goose droppings.     

Forage plants of an Arctic‐nesting herbivore show larger warming response in breeding than wintering grounds,potentially disrupting migration phenology

During spring migration, herbivorous waterfowl breeding in the Arctic depend on peaks in the supply of nitrogen‐rich forage plants, following a “green wave” of grass growth along their flyway to fuel migration and reproduction. The effects of climate warming on forage plant growth are expected to be larger at the Arctic breeding grounds than in temperate wintering grounds, potentially disrupting this green wave and causing waterfowl to mistime their arrival on the breeding grounds. We studied the potential effect of climate warming on timing of food peaks along the migratory flyway of the Russian population of barnacle geese using a warming experiment with open‐top chambers. We measured the effect of 1.0–1.7°C experimental warming on forage plant biomass and nitrogen concentration at three sites along the migratory flyway (temperate wintering site, temperate spring stopover site, and Arctic breeding site) during 2 months for two consecutive years. We found that experimental warming increased biomass accumulation and sped up the decline in nitrogen concentration of forage plants at the Arctic breeding site but not at temperate wintering and stop‐over sites. Increasing spring temperatures in the Arctic will thus shorten the food peak of nitrogen‐rich forage at the breeding grounds. Our results further suggest an advance of the local food peak in the Arctic under 1–2°C climate warming, which will likely cause migrating geese to mistime their arrival at the breeding grounds, particularly considering the Arctic warms faster than the temperate regions. The combination of a shorter food peak and mistimed arrival is likely to decrease goose reproductive success under climate warming by reducing growth and survival of goslings after hatching.     

Foraging behaviour and fuel accumulation of capital breeders during spring migration as derived from a combination of satellite‐ and ground‐based observations

The migration strategy of many capital breeders is to garner body stores along the flyway at distinct stopover sites. The rate at which they can fuel is likely to be strongly influenced by a range of factors, such as physiology, food availability, time available for foraging and perceived predation. We analysed the foraging behaviour and fuel accumulation of pink‐footed geese, an Arctic capital breeder, at their mid‐flyway spring stopover site and evaluated to what extent their behaviour and fuelling were related to physiological and external factors and how it differed from other stopovers along the flyway. We found that fuel accumulation rates of geese at the mid‐flyway site were limited by habitat availability rather than by digestive constraints. However, as the time available for foraging increased over the stopover season, geese were able to keep constant fuelling rate. Putting this in perspective, geese increased their daily net energy intake along the flyway corresponding to the increase in time available for foraging. The net energy intake per hour of foraging remained the same. Geese showed differences in their reaction to predators/disturbance between the sites, taking higher risks particularly at the final stopover site. Hence, perceived predation along the flyway may force birds to postpone the final fuel accumulation to the last stopover along the flyway. Flexibility in behaviour appears to be an important trait to ensure fitness in this capital breeder. Our findings are based on a new, improved method for estimating fuel accumulation of animals foraging in heterogeneous landscapes based on data obtained from satellite telemetry and habitat specific intake rates.     

Indirect effects of lemming cycles on sandpiper dynamics: 50 years of counts from southern Sweden

The bird-lemming hypothesis postulates that breeding success of tundra-nesting geese and waders in Siberia follows the cyclic pattern of lemming populations, as a result of predators switching from lemmings to birds when the lemming population crashes. We present 50 years of data on constant-effort catches of red knot Calidris canutus and curlew sandpiper C. ferruginea at an autumn migratory stopover site (Ottenby) at the Baltic Sea, supplemented with literature data on winter censuses of dark-bellied brent goose Branta b. bernicla and white-fronted goose Anser albifrons in northwestern Europe, and waders in Germany and Southern Africa. Number and proportion of juveniles in these bird populations (both our own and literature data) were compared with an index of predation pressure (calculated from the abundance of lemmings on the Taimyr peninsula), and climate indices for the North Eurasia and the North Atlantic regions. The index of predation pressure correlated significantly with the number of juveniles of red knot and curlew sandpiper, but not with number of adults. Also, this index correlated with the reproductive performance of geese and waders reported in the literature. Fourier analysis revealed a significant deviation from random noise with the maximum spectral density at the period length of 3 years for number of juvenile red knots and curlew sandpipers captured at Ottenby, abundance of lemmings, reproduction in arctic fox Alopex lagopus, and reproductive performance in geese on the Siberian tundra. Also, the date of passage at Ottenby for adult red knot and curlew sandpiper showed a spectral density peak at a period length of 3 years, the latter species also showing a peak at a period length of 5-6 years. Passage dates for adult red knot and curlew sandpiper were earlier in years of high predation pressure compared with years of low predation pressure. The fluctuations in reproductive success of the studied Siberian goose and wader species appear to be primarily influenced by biotic factors in the breeding area, rather than by abiotic factors, such as climate oscillations. Annual variations in migratory arctic bird populations may have far reaching effects in habitats along their migration routes and in their wintering areas. We suggest a link between lemming cyclicity in the Northern Hemisphere and predation pressure on Southern Hemisphere benthos, in which the signal is carried between continents by long distance migrating waders.     

基于卫星跟踪大天鹅春季迁徙时间及其影响因素

候鸟春季迁徙时间与其能否顺利完成迁徙过程,以及对繁殖地的成功选择和繁殖成效密切相关,通过对大天鹅越冬地和停歇地的春季迁徙时间选择原因及其影响因素进行分析,有助于深入理解候鸟春季迁徙时间策略和栖息地保护状况。2015年2月和12月,在河南三门峡湿地捕捉了60只越冬大天鹅并佩戴卫星跟踪器,获取了详细的大天鹅在越冬地和停歇地的春季迁徙时间等信息,并进一步分析了春季迁徙时间与气候因子的相关性。结果表明,大天鹅主要在夜间22：00-0：00和0：00-1：59迁离越冬地。大天鹅迁离越冬地的日期与温度呈显著性正相关,迁离时多选择顺风且风速较小的天气。大天鹅迁离越冬地后主要是在夜间飞行,而迁离停歇地后主要是在日间飞行。越冬地的温度越高,越有利于植物的生长,大天鹅可以快速地积累能量,提前开始春季迁徙。野外调查表明,内蒙古黄河中上游作为迁徙的重要停歇地,春季两岸捕鱼、农耕和放牧等为活动较多,因此大天鹅多选择在夜间觅食补充能量,在日间迁离。最后,针对黄河流域大天鹅栖息地的保护现状,提出了禁止经济开发项目、建立保护区和开展宣传教育等保护建议。     

Effects of agricultural change on abundance, fitness components and distribution of two arctic-nesting goose populations

Intensification of agriculture since the 1950s has enhanced the availability, competitive ability, crude protein content, digestibility and extended growing seasons of forage grasses. Spilled cereal grain also provides a rich food source in autumn and in winter. Long‐distance migratory herbivorous geese have rapidly exploited these feeding opportunities and most species have shown expansions in range and population size in the last 50 years. Results of long‐term studies are presented from two Arctic‐breeding populations, the Svalbard pink‐footed goose and the Greenland white‐fronted goose (GWFG). GWFGs have shown major habitat shifts since the 1950s from winter use of plant storage organs in natural wetlands to feeding on intensively managed farmland. Declines in local density on, and abandonment of, unmodified traditional wintering habitat and increased reproductive success among those birds wintering on farmland suggest that density‐dependent processes were not the cause of the shift in this winter‐site‐faithful population. Based on enhanced nutrient and energy intake rates, we argue that observed shifts in both species from traditionally used natural habitats to intensively managed farmland on spring staging and wintering areas have not necessarily been the result of habitat destruction. Increased food intake rates and potential demographic benefits resulting from shifts to highly profitable foraging opportunities on increasingly intensively managed farmland, more likely explain increases in goose numbers in these populations. The geographically exploratory behaviour of subdominant individuals enables the discovery and exploitation of new winter feeding opportunities and hence range expansion. Recent destruction of traditional habitats and declines in farming at northern latitudes present fresh challenges to the well being of both populations. More urgently, Canada geese colonizing breeding and moulting habitats of white‐fronted geese in Greenland are further affecting their reproductive output.     

Seagrass (Zostera spp.) as food for brent geese (Branta bernicla): an overview

Brent geese (called brant in North America) are among the smallest and the most marine of all goose species, and they have very long migration routes between high Arctic breeding grounds and temperate wintering grounds. Like all other geese, brent geese are almost entirely herbivorous. Because of these ecological characteristics they have a high food demand and are strongly dependent on stopover sites to ”refuel” during the migration period. Three subspecies of brent geese are distributed around the Holarctic, forming seven populations with distinct migration routes. Most or all of these populations make heavy use of Zostera spp. during migratory stopovers on spring and/or autumn migration. Examples of Zostera stopover areas being used by large numbers of brent geese for several weeks each year are Izembek Lagoon (Alaska), lagoons in Baja California, the German/Danish Wadden Sea, the Golfe du Morbihan (France), British estuaries, and the White Sea (Western Russian Arctic). Brent geese feed on Zostera wherever they can, but they can only reach the plants at low tide or in shallow water. Changes in Zostera abundance affect brent goose distribution, and the ”wasting disease” affecting Atlantic Zostera stocks during the 1930s was at least partly responsible for a steep decline in brent goose population sizes on both sides of the Atlantic. While Zostera is of outstanding importance as food for brent geese, the impact of the geese on Zostera stocks seems to be less important – at many sites, the geese consume only a small amount of the available Zostera, or, if they consume more, the seagrass can regenerate fully until the following season. Received: 6 December 1998 / Received in revised form: 6 August 1999 / Accepted: 9 August 1999     

Light‐level geolocation reveals wintering distribution,migration routes,and primary stopover locations of an endangered long‐distance migratory songbird

The importance of understanding the geographic distribution of the full annual cycle of migratory birds has been increasingly highlighted over the past several decades. However, the difficulty of tracking small birds between breeding and wintering areas has hindered progress in this area. To learn more about Kirtland's warbler Setophaga kirtlandii movement patterns throughout the annual cycle, we deployed archival light‐level geolocators across their breeding range in Michigan. We recovered devices from 27 males and analyzed light‐level data within a Bayesian framework. We found that most males wintered in the central Bahamas and exhibited a loop migration pattern. In both fall and spring, departure date was the strongest predictor of arrival date, but in spring, stopover duration and migration distance were also important. Though stopover strategies varied, males spent the majority of their spring migration at stopover sites, several of which were located just before or after large ecological barriers. We argue that loop migration is likely a response to seasonal variation in prevailing winds. By documenting a tight link between spring departure and arrival dates, we provide a plausible mechanism for previously documented carry‐over effects of winter rainfall on reproductive success in this species. The migratory periods remain the least understood periods for all birds, but by describing Kirtland's warbler migration routes and timing, and identifying locations of stopover sites, we have begun the process of better understanding the dynamics of their full annual cycle. Moreover, we have provided managers with valuable information on which to base future conservation and research priorities.     

Spring migratory routes and stopover duration of satellite‐tracked Eurasian Teals Anas crecca wintering in Italy

Identifying an organism's migratory strategies and routes has important implications for conservation. For most species of European ducks, information on the general course of migration, revealed by ringing recoveries, is available, whereas tracking data on migratory movements are limited to the largest species. In the present paper, we report the results of a tracking study on 29 Eurasian Teals, the smallest European duck, captured during the wintering period at three Italian sites. The departure date of spring migration was determined for 21 individuals, and for 15 the entire spring migratory route was reconstructed. Most ducks departed from wintering grounds between mid‐February and March following straight and direct routes along the Black Sea‐Mediterranean flyway. The breeding sites, usually reached by May, were spread from central to north‐Eastern Europe to east of the Urals. The migratory speed was slow (approximately 36 km/day on average) because most birds stopped for several weeks at stopover sites, mainly in south‐eastern Europe, especially at the very beginning of migration. The active flight migration segments were covered at much higher speeds, up to 872 km/day. Stopover duration tended to be shorter when birds were closer to their breeding site. These results, based on the largest satellite tracking effort for this species, revealed for the first time the main features of the migratory strategies of individual Teals wintering in Europe, such as the migration timing and speed and stopover localization and duration.     

Potential for an Arctic‐breeding migratory bird to adjust spring migration phenology to Arctic amplification

Arctic amplification, the accelerated climate warming in the polar regions, is causing a more rapid advancement of the onset of spring in the Arctic than in temperate regions. Consequently, the arrival of many migratory birds in the Arctic is thought to become increasingly mismatched with the onset of local spring, consequently reducing individual fitness and potentially even population levels. We used a dynamic state variable model to study whether Arctic long‐distance migrants can advance their migratory schedules under climate warming scenarios which include Arctic amplification, and whether such an advancement is constrained by fuel accumulation or the ability to anticipate climatic changes. Our model predicts that barnacle geese Branta leucopsis suffer from considerably reduced reproductive success with increasing Arctic amplification through mistimed arrival, when they cannot anticipate a more rapid progress of Arctic spring from their wintering grounds. When geese are able to anticipate a more rapid progress of Arctic spring, they are predicted to advance their spring arrival under Arctic amplification up to 44 days without any reproductive costs in terms of optimal condition or timing of breeding. Negative effects of mistimed arrival on reproduction are predicted to be somewhat mitigated by increasing summer length under warming in the Arctic, as late arriving geese can still breed successfully. We conclude that adaptation to Arctic amplification may rather be constrained by the (un)predictability of changes in the Arctic spring than by the time available for fuel accumulation. Social migrants like geese tend to have a high behavioural plasticity regarding stopover site choice and migration schedule, giving them the potential to adapt to future climate changes on their flyway.     

Interactions between land use,habitat use,and population increase in greater snow geese: what are the consequences for natural wetlands?

The North American greater snow goose population has increased dramatically during the last 40 years. We evaluated whether refuge creation, changes in land use on the wintering and staging grounds, and climate warming have contributed to this expansion by affecting the distribution, habitat use, body condition, and migration phenology of birds. We also reviewed the effects of the increasing population on marshes on the wintering grounds, along the migratory routes and on the tundra in summer. Refuges established before 1970 may have contributed to the initial demographic increase. The most important change, however, was the switch from a diet entirely based on marsh plants in spring and winter (rhizomes of Scirpus/Spartina) to one dominated by crops (corn/young grass shoots) during the 1970s and 1980s. Geese now winter further north along the US Atlantic coast, leading to reduced hunting mortality. Their migratory routes now include portions of southwestern Québec where corn production has increased exponentially. Since the mid‐1960s, average temperatures have increased by 1–2.4°C throughout the geographic range of geese, which may have contributed to the northward shift in wintering range and an earlier migration in spring. Access to spilled corn in spring improved fat reserves upon departure for the Arctic and may have contributed to a high fecundity. The population increase has led to intense grazing of natural wetlands used by geese although these habitats are still largely undamaged. The foraging in fields allowed the population to exceed limits imposed by natural marshes in winter and spring, but also prevented permanent damage because of their overgrazing.     

Seasonal migrations of four individual bar-headed geese Anser indicus from Kyrgyzstan followed by satellite telemetry

The Kyrgyz population of the bar-headed goose Anser indicus has declined dramatically during the past decades. Human persecution during migration and habitat loss at stopover and wintering sites are commonly regarded as most serious threats. However, little is known about seasonal movements, migration routes, and wintering sites of the bar-headed geese from Kyrgyzstan, which represent the westernmost geographical population of the species. As part of a conservation project, which also included reinforcement of the wild population by the release of hand-reared juveniles, in late summer of 1998, five bar-headed geese, three wild adults and two hand-reared goslings, were fitted with sun-powered satellite transmitters in order to track their movements from Lake Son Kul and Lake Chatyr Kul in Kyrgyzstan. The five individuals contributed very unevenly to the more than 5,000 signals in total that were received from the French ARGOS system: one failed after 8 weeks, while another one was tracked for more than 2 years. The four geese contributing to this study followed three completely different migration routes leading to their wintering areas in Pakistan, India and Uzbekistan, while stopover areas were situated in southern Tajikistan and in western Tibet. Both in autumn and spring the adult birds migrated distances of 1,280–1,550 km in two steps, with stopover periods of 32–46 days (autumn) and 16–23 days (spring). Flight speeds of up to 680 km per actual migration day were recorded regularly, even during crossings of very high summits. A hand-reared juvenile flew non-stop for 790 km to southern Uzbekistan and even visited southernmost Turkmenistan, where the species is very rarely seen. The timing of migration varied considerably between individuals but also for the same individual between years. We compare our tracking results with previous findings (field observations, ring recoveries, and satellite tracking results) and discuss them with respect to migration over high-mountain habitats and a general migration strategy of the species.     

Faster migration in autumn than in spring: seasonal migration patterns and non‐breeding distribution of Icelandic whimbrels Numenius phaeopus islandicus

Migration is fundamental in the life of many birds and entails significant energetic and time investments. Given the importance of arrival time in the breeding area and the relatively short period available to reproduce (particularly at high latitudes), it is expected that birds reduce spring migration duration to a greater extent than autumn migration, assuming that pressure to arrive into the wintering area might be relaxed. This has previously been shown for several avian groups, but recent evidence from four tracked Icelandic whimbrels Numenius phaeopus islandicus, a long distance migratory wader, suggests that this subspecies tends to migrate faster in autumn than in spring. Here, we 1) investigate differences in seasonal migration duration, migration speed and ground speed of whimbrels using 56 migrations from 19 individuals tracked with geolocators and 2) map the migration routes, wintering and stopover areas for this population. Tracking methods only provide temporal information on the migration period between departure and arrival. However, migration starts with the fuelling that takes place ahead of departure. Here we estimate the period of first fuelling using published fuel deposition rates and thus explore migration speed using tracking data. We found that migration duration was shorter in autumn than in spring. Migration speed was higher in autumn, with all individuals undertaking a direct flight to the wintering areas, while in spring most made a stopover. Wind patterns could drive whimbrels to stop in spring, but be more favourable during autumn migration and allow a direct flight. Additionally, the stopover might allow the appraisal of weather conditions closer to the breeding areas and/or improve body condition in order to arrive at the breeding sites with reserves.     

Could Have Gone Wrong: Effects of Abrupt Changes in Migratory Behaviour on Harvest in a Waterbird Population

To sustainably exploit a population, it is crucial to understand and reduce uncertainties about population processes and effects of harvest. In migratory species, management is challenged by geographically separated changing environmental conditions, which may cause unexpected changes in species distribution and harvest. We describe the development in the harvest of Svalbard-breeding pink-footed geese (Anser brachyrhynchus) in relation to the observed trajectory and migratory behaviour of the population. In autumn, geese migrate via stopover sites in Norway and Denmark (where they are hunted) to wintering grounds in the Netherlands and Belgium (where they are protected). In Denmark and Norway harvesting increased stepwise during the 2000s. The increase in the population size only partly explained the change. The change corresponded to a simultaneous stepwise increase in numbers of geese staging in Denmark throughout autumn and winter; geese also moved further inland to feed which collectively increased their exposure to hunting. In Norway the increase in harvest reflected greater utilisation of lowland farmland areas by geese, increasing their hunting exposure. The study demonstrates how changes in migratory behaviour can abruptly affect exposure to hunting, which showed a functional response to increased temporal and spatial availability of geese. The harvest has now reached a level likely to cause a population decline. It highlights the need for flexible, internationally coordinated hunting regulations and reliable up-to-date population estimates and hunting bag statistics, which are rare in European management of migratory waterbirds. Without such information decisions are left with judgments based on population estimates, which often have time lags of several years between recording and reporting, hampering possibilities for the timely adjustment of management actions.     

Reinterpretation of gizzard sizes of red knots world-wide emphasises overriding importance of prey quality at migratory stopover sites

The size of digestive organs can be rapidly and reversibly adjusted to ecological circumstances, but such phenotypic flexibility comes at a cost. Here, we test how the gizzard mass of a long-distance migrant, the red knot (Calidris canutus), is adjusted to (i) local climate, (ii) prey quality and (iii) migratory fuelling demands. For eight sites around the world (both wintering and stopover sites), we assembled data on gizzard masses of free-living red knots, the quality of their prey and the local climate. Using an energetic cost-benefit approach, we predicted the gizzard size required for fastest fuelling (net rate-maximization, i.e. expected during migration) and the gizzard size required to balance daily energy budgets (satisficing, expected in wintering birds) at each site. The measured gizzards matched the net rate-maximizing predictions at stopover sites and the satisficing predictions at wintering sites. To our surprise, owing to the fact that red knots selected stopover sites with prey of particularly high quality, gizzard sizes at stopovers and at wintering sites were nevertheless similar. To quantify the benefit of minimizing size changes in the gizzard, we constructed a model incorporating the size-dependent energy costs of maintaining and carrying a gizzard. The model showed that by selecting stopovers containing high-quality prey, metabolic rates are kept at a minimum, potentially reducing the spring migratory period by a full week. By inference, red knots appear to time their stopovers so that they hit local peaks in prey quality, which occur during the reproductive seasons of the intertidal benthic invertebrates.