Differences in shifts of wintering and breeding ranges lead to changing migration distances in European birds

Studies on the impact of climate change on the distributions of bird species in Europe have largely focused on one season at a time, especially concerning summer breeding ranges. We investigated whether migratory bird species show consistent range shifts over the past 55 yr in both breeding and wintering areas or if these shifts are independent. We then analyzed whether patterns in changing migration distances of Finnish breeding birds could be explained by habitat use, phylogeny or body size. We used long‐term datasets from the Finnish ringing centre to analyze the mean wintering latitudes of 29 species of Finnish breeding birds, then used breeding distribution data to make predictions as to whether certain species were migrating shorter or longer distances based on the comparative shifts in the wintering and breeding grounds. Our data reveal species‐specific differences in changing migration distances. We show that for many species, long‐term shifts in wintering ranges have not followed the same patterns as those in the breeding range, leading to differences in migration distances over time. We conclude that species are not adjusting predictably to climate change in their wintering grounds, leading to changing migration distances in some, but not all, species breeding in Finland. This research fills an important gap in the current climate change biology literature, focusing on individuals’ entire life histories and revealing new complexities in range shift patterns.     

Climate determinants of breeding and wintering ranges of lesser kestrels in Italy and predicted impacts of climate change

Climate warming would theoretically create conditions for the breeding range expansion of pseudo‐steppe Mediterranean and long‐distance migrant species and provide the possibility for these to overwinter in the same breeding areas. However, contemporary changes in rainfall regimes might have negative effects on the climate suitability and in turn, shrink species potential range. The lesser kestrel Falco naumanni is highly sensitive to rainfall oscillations and has recently extended its Italian breeding range towards northern latitudes and increasing its wintering records. We modelled the effects of temperature and rainfall on current and future climate suitability for lesser kestrels in both the breeding and wintering periods by using MaxEnt. Models were based on the distribution of 298 colonies and 40 wintering records. Future climate suitability was assessed under eight different scenarios. Spring rainfall amount resulted as the main determinant of breeding climate suitability, so its predicted reduction will determine a shrinkage in suitable areas (–42.10% in 2050; –32.07% in 2070). Specifically, the 66.05% of Italian colonies will be outside the climatically suitable area by 2050. However wide areas, suitable under current climate conditions, are still not occupied by lesser kestrel and allow the potential expansion of its Italian breeding range in the short term. Temperature seasonality mainly determined the species’ winter climate suitability, which is overall predicted to boost in the next decades (+145.03% in 2050; and +123.91% in 2070). All but one future scenarios predicted a northward shift of about 40 km for both breeding and wintering climate suitability. Despite its recent expansion, we have found that climate change will pose conservation concerns for the Italian breeding population of lesser kestrels. Indeed, changes in non‐climate factors will also outline the future suitability of the Italian range for lesser kestrels in both seasons with effects that might both strengthen or mitigate climate effects.     

Understanding population change in migratory songbirds: long-term and experimental studies of Neotropical migrants in breeding and wintering areas

Effective conservation and management of migratory bird species requires an understanding of when and how their populations are limited and regulated. Since 1969, my colleagues and I have been studying migratory songbird populations in their breeding quarters at the Hubbard Brook Experimental Forest in north-central New Hampshire, USA, and since 1986, in their winter quarters in the Greater Antilles (Jamaica). Long-term data on the abundance and demography of these populations, coupled with experimental tests of mechanisms, indicate that processes operating in the breeding area (e.g. density-dependent fecundity, food limitation) are sufficient to limit and regulate the local abundance of these species. At the same time, limiting factors operating in the non-breeding season (e.g. climate-induced food limitation in winter quarters and especially mortality during migration) also have important impacts on migrant populations. Furthermore, recent studies have shown that limiting processes during the winter period can carry over into the breeding season and affect reproductive output. These findings clearly demonstrate that to understand changes in abundance of long-distance migrant species requires knowledge of events operating throughout the annual cycle, which presents a challenge to researchers, managers and others concerned with the welfare of these species.     

Understanding population change in migratory songbirds: long‐term and experimental studies of Neotropical migrants in breeding and wintering areas

Effective conservation and management of migratory bird species requires an understanding of when and how their populations are limited and regulated. Since 1969, my colleagues and I have been studying migratory songbird populations in their breeding quarters at the Hubbard Brook Experimental Forest in north‐central New Hampshire, USA, and since 1986, in their winter quarters in the Greater Antilles (Jamaica). Long‐term data on the abundance and demography of these populations, coupled with experimental tests of mechanisms, indicate that processes operating in the breeding area (e.g. density‐dependent fecundity, food limitation) are sufficient to limit and regulate the local abundance of these species. At the same time, limiting factors operating in the non‐breeding season (e.g. climate‐induced food limitation in winter quarters and especially mortality during migration) also have important impacts on migrant populations. Furthermore, recent studies have shown that limiting processes during the winter period can carry over into the breeding season and affect reproductive output. These findings clearly demonstrate that to understand changes in abundance of long‐distance migrant species requires knowledge of events operating throughout the annual cycle, which presents a challenge to researchers, managers and others concerned with the welfare of these species.     

Behavioral and physiological conflicts in migrants: the transition between migration and breeding

Progression of the vernal migratory life history stage to breeding presents a number of apparent behavioral and physiological conflicts. Features that characterize the migratory stage include: high mobility, sociality, repetitive cycles of feeding (hyperphagia and fattening) and migratory flight. Breeding comprises: sedentary, territorial and reproductive behaviors, an initial decline in hyperphagia and reduction of fuel stores. Because morphology, physiology and behavior change, the transition between stages cannot be instantaneous. In many species development of the reproductive system actually occurs during migration, but in others gonadal development may not commence until later. This variation in degree of overlap of migration and reproductive functions is not well understood, but may be related to migratory distance and length of the breeding season, which tends to be shorter at higher latitudes and altitudes. In these habitats, migrants may arrive at their breeding sites to find unpredictable conditions that cannot support breeding. At this juncture, migrants may retreat to refugia and prolong maintenance of facultative migratory functions, termed arrival biology, until conditions improve sufficiently to initiate breeding. In this review, we focus on the Pacific races of the white-crowned sparrow, Zonototrichia leucophyrs, in which the entire spectrum of migratory strategies are represented from resident to long distance migrants and about which much is known. This species presents a unique view of the appearance and variations in arrival biology. Focusing on the juncture between migration and breeding, we discuss the diversity of responses of congeners to a spectrum of environmental conditions that favor survival and reproductive success.     

Assigning birds to wintering and breeding grounds using stable isotopes: lessons from two feather generations among three intercontinental migrants

Geographic origins of populations and migration patterns of several vertebrate and invertebrate species have been inferred from geographically distinct isotopes in their tissues. To test the hypothesis that feathers grown on different continents would reflect continental differences of δD in precipitation and have significantly different stable isotope ratios, we analyzed stable isotopes in two generations of feathers from three bird species (American and Pacific golden-plovers, Pluvialis dominica and P. fulva, and northern wheatears Oenanthe oenanthe) that breed in North America and winter in South America, the South Pacific and Asia, and Africa. We found significant differences in stable isotope signatures between summer- and winter-grown feathers in the plovers, and our use of two generations of feathers provided similar variation to that reported in studies using larger sample sizes. In contrast to plovers, no differences were detected in isotope values between summer- and winter-grown feathers in wheatears. Discriminant analyses separated 80% of summer- and winter-grown feathers for each plover species. Nonetheless, an “assignment with exclusion” method adapted from population genetics to impart a measure of confidence in assigning individuals to groups of origin resulted in an overall accuracy among plovers of only 41%, compared with a 63% assignment accuracy when the exclusion criterion was removed. Thus, we were unable to accurately assign feathers to origin of growth on the continental scale. Moreover, using δD expectations for North America, we were unable to assign summer-grown plover feathers to within better than several thousand kilometers of their true origins. We urge researchers to carefully consider the ecology and physiology of their study organisms, statistical methodology, and the interpretation of results when using stable isotopes to infer the geographic origins of feather growth.     

Should females of migratory dabbling ducks switch mates between wintering and breeding sites?

Pairing of northern hemisphere dabbling ducks normally occurs in wintering sites. Mate choice occurs when some individuals congregate in courtship parties, in which females mainly choose mates according to male behavioural dominance. The sex ratio of some species is more male-biased at northern than at southern wintering sites, and the age ratio is adult-biased in northern areas. A possible mechanism responsible for this spatial segregation of sex and age classes is behavioural dominance, with males usually dominating females and adults dominating yearlings. Due to this latitudinal segregation of sexes during winter, presumably more males would remain unpaired in northern sites utilized by the more dominant birds than in southern sites utilized by the less-dominant birds, and this is paradoxical. I argue that some females having wintered and paired in southern sites may switch mates at stopover sites during northward migration. This hypothesis provides an explanation as to why, in spite of females being paired when initiating northward migration, some species show an increase in courtship activity during spring at stopover sites, as in common teals (Anas crecca) in southern Spain. In contrast, populations that do not migrate, e.g. mallards (A. platyrhynchos) in southern Spain, do not exhibit an increase in courtship activity during spring.     

Conservation of migrants on their wintering grounds: an overview

Most research on the behaviour and ecology of wintering migrants has been undertaken in the Neotropics. In Africa, conservation biologists are faced with a comparative paucity of baseline data to assess the significance of environmental change, in contrast to the comparatively well-developed monitoring programmes in place on the breeding grounds. A programme of monitoring, established through international collaboration and matched with further ecological research, is required. Given the 'dispersed' distribution of most passerine migrants, determination of the significance of apparent detrimental effects can be hard to ascertain. However, migrants may provide conservationists with an effective vehicle for the provision of input into region-wide development policy. It is only through influencing regional development policies within the continent that large-scale and long-term conservation of important bird habitats will be achieved.     

Geolocators reveal migratory connectivity between wintering and breeding areas of Golden‐winged Warblers

The conservation of migratory songbirds is often impeded by a lack of understanding of how populations in breeding and wintering areas are geographically linked (migratory connectivity). In recent years, light‐level geolocators have improved our understanding of migratory connectivity. Such information is valuable for evaluating how conservation efforts align between the breeding and non‐breeding areas of at‐risk species, and help to more effectively prioritize the allocation of conservation funding. Golden‐winged Warblers (Vermivora chrysoptera) are imperiled migratory songbirds, but the extent to which conservation efforts in their breeding and non‐breeding areas coincide with patterns of migratory connectivity are not well known. We used light‐level geolocators to evaluate the extent to which conservation actions targeting Golden‐winged Warblers in Nicaragua and in their breeding range in North America align with patterns of migratory connectivity. We recovered six of 22 geolocators that had been deployed on male Golden‐winged Warblers at the El Jaguar Reserve during the winter of 2015–2016. All six males migrated to breeding areas in the western Great Lakes region that includes eastern Minnesota, northern Wisconsin, southwestern Ontario, and Michigan's Upper Peninsula. All six males also had similar migration routes, with spring stopovers in southern Mexico, Guatemala, and Belize, a trans‐Gulf flight, and a stopover in the region of Louisiana, Arkansas, eastern Oklahoma, and Texas. Our results, in combination with those of previous studies, demonstrate strong migratory connectivity between portions of the breeding and winter distributions of Golden‐winged Warblers currently targeted for conservation. However, additional studies are needed to improve our understanding of the stopover ecology of Golden‐winged Warblers, especially in areas where they remain for extended periods of time. Finally, patterns of migratory connectivity revealed in our study should be used in combination with existing demographic parameters for Golden‐winged Warblers in the western Great Lakes and Nicaragua to help inform full life cycle population models for this imperiled songbird.     

春化作用与甘蓝类蔬菜的生育障碍(综述)

本文着重探讨结球甘蓝未熟抽薹和花椰菜花球劣变的发生与春化作用的内在联系,系统介绍结球甘蓝和花椰菜的春化特性及春化条件,详细分析其生育障碍产生的原因,并提出合理的预防措施,从而解决结球甘蓝和花椰菜生产栽培中的难题。     

Conservation of migrants on their breeding grounds

This paper addresses the question of how to diagnose the causes of a population decline of a migratory bird so that conservation action can be taken. Diagnosis may be through study of habitats or population factors. Different approaches are illustrated with examples from the more threatened migratory species which breed in Europe. Gaps in knowledge of how to make conservation diagnoses and of the circumstances of some of Europe's few migratory and globally endangered birds are shown.     

Population limitation in Palaearctic-African migrant passerines

We report the results of key factor analyses for Sedge Warbler Acrocephalus schoenobaenus, Blackcap Sylvia atricapilla, Whitethroat Sylvia communis and Willow Warbler Pftylloscopus Irochilus based on extensive data from the United Kingdom gathered by the British Trust for Ornithology. These results are compared with those from published key factor analyses based on intensive studies of Swallows Hirundo ruslica, Redstarts Phoenicurus phoenicums and three populations of Pied Flycatchers Fiiedula hypoleuca. Variation in losses between fledging and the following breeding season lk₄ ) accounted for most of the variation in total losses (K_T) for all populations. Breeding performance was poorly correlated with population changes. Density-dependence was found mainly in 6₄, its strength decreasing as average rate of population change increased. There was also evidence of weak density-dependence in the clutch size of Swallows and in Willow Warbler egg losses. Fluctuations in k₄ were correlated with conditions on the wintering grounds in Sedge Warblers, Whitethroats and Swallows, for the first two species these relationships being confirmed by analyses of independent survival estimates based on mark-recapture data. Populations of Sedge Warblers and Whitethroats appear to be limited by competition for resources on the wintering grounds. Removal experiments suggest that Willow Warblers, Pied Flycatchers and perhaps Blackcaps are limited through recruitment to the breeding population, in the case of Pied Flycatchers this limitation being in relation to the availability of nest sites.     

The importance of Prespa National Park for breeding and wintering birds

The breeding avifauna of Prespa National Park is ofnational and international importance both due to itshigh richness and the internationally importantpopulations of rare species. The latter include theDalmatian (Pelecanus crispus) and Great White(Pelecanus onocrotalus) pelicans and the PygmyCormorant (Phalacrocorax pygmaeus). Populationsof national interest include Cormorant (Phalacrocorax carbo), Greylag Goose (Anseranser), Goosander (Mergus merganser), GreatWhite Egret (Egretta alba) and six otherspecies of ardeids. Among the 261 bird speciesobserved in the area of ca 250 km² since thelate 60s, 164 breed and 41 are very rarely observed.Passage migrants and winter visitors include 91species. Twelve breeding species became extinct in theNational Park during the last 25 years and others areendangered. Raptors and aquatic birds suffered moredue to habitat and land use alterations brought aboutby the construction of an irrigation scheme.Eighty-one species observed in the area are listed asin need of special protection in Annex I of the 79/409EEC Directive and 32 of them breed. Lake Mikri Prespais of virtually no importance for wintering waterfowlsince it is frozen every winter, but the Greek part ofLake Megali Prespa, which never freezes, concentratesamong other species the 20% and 35% respectively ofthe Great Crested Grebes and Tufted Ducks wintering inGreece.     

Migration routes and wintering areas of Booted Eagles Aquila pennata breeding in Spain

Capsule?Five Booted Eagles breeding in Spain were tracked by GPS during migration. Autumn routes were generally more eastern than spring routes, showing a typical loop migration. Birds covered on average ca. 200 km/day, and only one individual used a long-term stopover site (for up to 4 weeks). All but one used a single wintering area, located in Sub-Saharan Africa, at 2800–3500 km from their nests. Eagles were forced to stop migration at the Strait of Gibraltar for up to 6 days.     

Strategies, punctuality of arrival and ranges of migrants in the Kalahari basin, Botswana

The timing and intensity of the first summer rains in the Kalahari basin are highly variable between years. Avian migrants to the area follow two principal strategies. Species arriving before November typically do so before significant rain can be expected and do not use ecological conditions as a proximal cue to time arrival; their arrival was unaffected by the failure of the early rains in 1994. Species arriving after mid-November respond to the timing and extent of the early rains and were progressively more delayed in parallel to a cumulative deficit in rainfall during the early part of the 1994 wet season. The second strategy results in erratic occupation of parts of the nonbreeding range, so that during drought years, rain-dependent, late-arriving species may remain absent from the Kalahari basin, complicating the assessment of the "effective" size of their nonbreeding range.     

Linking the wintering and breeding grounds of warblers along the Pacific Flyway

Long‐distance migration is a behavior that is exhibited by many animal groups. The evolution of novel migration routes can play an important role in range expansions, ecological interactions, and speciation. New migration routes may evolve in response to selection in favor of reducing distance between breeding and wintering areas, or avoiding navigational barriers. Many migratory changes are likely to evolve gradually and are therefore difficult to study. Here, we attempt to connect breeding and wintering populations of myrtle warblers (Setophaga coronata coronata) to better understand the possible evolution of distinct migration routes within this species. Myrtle warblers, unlike most other warblers with breeding ranges primarily in eastern North America, have two disjunct overwintering concentrations—one in the southeastern USA and one along the Pacific Coast—and presumably distinct routes to‐and‐from these locations. We studied both myrtle and Audubon's warblers (S. c. auduboni) captured during their spring migration along the Pacific Coast, south of the narrow region where these two taxa hybridize. Using stable hydrogen isotopes and biometric data, we show that those myrtle warblers wintering along the southern Pacific Coast of North America are likely to breed at high latitudes in Alaska and the Yukon rather than in Alberta or further east. Our interpretation is that the evolution of this wintering range and migration route along the Pacific Coast may have facilitated the breeding expansion of myrtle warblers into northwestern North America. Moreover, these data suggest that there may be a migratory divide within genetically similar populations of myrtle warblers.     

Abundance-range size relationships of breeding and wintering birds in Britain: a comparative analysis

Both breeding and wintering assemblages of birds in Britain exhibit positive interspecific relationships between population size and geographic range size, such that the average density of species is greater if they are more widely distributed Species in common to both assemblages, that is resident species, had greater population sizes, geographic range sizes, and densities in winter In contrast, whilst winter migrants had higher abundances than summer migrants, the range sizes of the former were disproportionately larger still, resulting in a lower density for species that only winter in Britain than for those that only breed Such differences aside, the overall form of the abundance-range size relationship is remarkably similar between the two assemblages and their constituent subsets of species     

Migration patterns, breeding and moulting locations of king eiders wintering in north-eastern Norway

The Norwegian coast is a very important winter area for king eiders (Somateria spectabilis), but their origin has been unknown. We determined spring and autumn migration routes, timing of migration and potential breeding areas of king eiders wintering in north-eastern Norway using implanted satellite transmitters. Five males and five females were equipped with transmitters in February 2008, and location data were received from six birds. All birds departed within 10 days in mid April and flew to the Pechora Sea and Kara Sea in western Russia where they staged until mid June. Subsequently, four of five birds with active transmitters (two females and two males) moved east to potential breeding locations on the Gydan and Taymyr Peninsulas. In early July, the males moved west to moult at Kolguyev Island and in the Pechora Sea. The two females departed in mid July, one probably moulting between the gulfs of the Ob and Yenisei Rivers, after which it moved to the Pechora Sea. The other female probably moulted in the eastern Taymyr, thereafter moving to Novaya Zemlya. This female returned to the north-eastern coast of Norway 1 December, while the other female returned 2 January. For the males, the transmitters stopped in December/January when they were still in the Pechora/Kolguyev area. King eiders wintering in north-eastern Norway thus originate from the western half of Arctic Russia, and the Taymyr Peninsula is probably the dividing point between the eastern and western flyways.