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Transcontinental pathways and seasonal movements of an Asian migrant, the Common Ringed Plover Charadrius hiaticula tundrae
Abstract
Little is known about the migration of Common Ringed Plovers Charadrius hiaticula in the Asian part of their range. This study describes the seasonal movements of birds from the easternmost population of the subspecies tundrae (S Chukotka, E Russia) based on data from geolocators. The wintering grounds of five males were scattered from the Persian Gulf to the Nile Delta and south to Somalia. During the wintering period three birds made only local movements, whereas two others moved 1,100 and 3,200 km northward in the second half of March, before embarking on pre-breeding migration one month later. During post-breeding migration, the birds used a ‘hopping’ strategy, following the inland West Asian–East African Flyway and making hundreds of stops of various lengths, as recorded by their geolocators’ conductivity sensors. We recognize three main regional stopover areas during southward and northward migration. The pathways of each bird were rather similar in autumn and spring in the southwestern half of the routes, while a loop migration was evident in Siberia, and although the distances covered were longer, the migration speed was faster on the northbound route than on the southbound one. The migration tracks likely reflect a historical eastward expansion of the breeding grounds of this species.
Supplementary resources (2)
Data
February 2018
Data
February 2018
... psammodromus necessarily includes nonstop flights of at least 800-2,000 km across the North Atlantic, and the most northeasterly breeders of C. h. tundrae travel oneway distances of 9,000-13,000 km (Tomkovich et al. 2017), and perhaps more than 16,000 km to southern Africa (Underhill et al. 1999), which would be the longest migration of any plover. ...
... One example is the Common Ringed Plover, in which migration to the relatively constricted nonbreeding range involves an eastward movement of ≥73° of longitude for the westernmost breeders and ≥100° westward for the easternmost breeders (Figure 7.1). The extreme example is an individual tracked from Chukotka to the Nile River delta, a longitudinal shift of 146° (Tomkovich et al. 2017). Similarly, all Eurasian Dotterels spend the winter in a narrow temperate band from Morocco to the Red Sea, despite a breeding range that spans from western Europe across the Palearctic and into western Alaska. ...
... Until recently, no individuals in this group had been remotely tracked on migration, largely because lightweight and long-lived tracking devices suitable for migratory birds weighing <80 g have only recently become available (Bridge et al. 2011). Three Charadrius species have now been tracked using miniaturized (~1 g) light-level geolocators, providing the first insights into year-round individual migration strategies (Minton et al. 2011, Hedenström et al. 2013, Lislevand et al. 2017, Tomkovich et al. 2017. Such studies will surely become more common in the near future, as tracking devices continue to decrease in size and cost, and methods for analyzing tracking data continue to improve. ...
Plovers of the subfamily Charadriinae demonstrate nearly every conceivable annual movement pattern found among birds, including long- and short-distance latitudinal migration, altitudinal migration, irruption, nomadism, and sedentary habits. Within species, and even within some populations, inter-individual differences may range from completely sedentary birds to some of the longest avian migrations yet recorded. This extreme variation makes plovers an interesting group in which to explore the ecological and evolutionary drivers and consequences of movement patterns. In this chapter, I review patterns of movement according to geography, molt, wing morphology, various aspects of annual-cycle strategies, and the known evolutionary history of the group. Of 40 plover species, 26 (65%) are migratory to some degree, and these are found on nearly all global migratory flyways, with the greatest number on the East Asian-Australasian Flyway. The species diversity and the proportion of non-migratory plovers are highest in the Eastern and Southern Hemispheres. Migration distance increases with breeding latitude; the longest migrations are performed by Arctic-breeding plovers, and no Southern Hemisphere breeders migrate further than the Tropics. In general, individual migration strategies are poorly described, but include both short-hop migrations and non-stop flights of at least 5,300 km and perhaps more than 7,000 km in some species. Wing shape varies with migration distance, suggesting that movement patterns and morphology co-evolve to some degree, and that some species may be pre-disposed to greater flexibility in movements. Repeated loss of migration appears to be a significant form of diversification in plovers; recent phylogenetic evidence supports historical radiation from a northern migratory ancestor, and current distributions of sedentary species, particularly on islands, suggests a pattern of isolation from mainland migratory species. Although some species show evidence of evolutionary constraints on migration routes, the present-day diversity of movement patterns implies great flexibility to respond to changing circumstances at multiple time scales. However, the general lack of specific information regarding routes and habitats used during migration is a major obstacle to developing effective global conservation strategies for migratory plovers.
... The global population is estimated at 360,000-1.3 million individuals (Delany et al. 2006), and Iceland holds the largest European population (up to 50,000 breeding pairs; Wiersma et al. 2018). Return rates of juveniles to their hatching site is relatively low, implying high natal dispersal, whereas adult return rates to breeding sites seem to be variable (Laven 1940, Wallander & Andersson 2003, Lislevand et al. 2017, Tomkovich et al. 2017. Western Palearctic Ringed Plovers show leap-frog migration, with the northern breeding populations generally wintering further south than southern populations (Taylor 1980). ...
... Interestingly, plovers recently started to close this gap in the breeding distribution using artificial gravel or sand mounds created by humans near villages, suggesting that the colonisation process is still ongoing (Lappo et al. 2012). The current migration routes of Ringed Plovers breeding in Chukotka support the idea of their postglacial eastward expansion: Chukotka Ringed Plovers have been found to winter in Africa and the Middle East, but during both spring and autumn migration, they migrate through Siberia following the West Asian -East African Flyway (Tomkovich et al. 2017). ...
... Such high juvenile dispersal will result in high gene flow and prevent population divergence since as little as one migrant per generation is commonly assumed to prevent populations from diverging (Mills & Allendorf 1996). Recent studies have also pointed to individual variation in migratory behaviour within subspecies with variation in routes, periods and selection of stop over sites, leading to shorter 'hops' and longer 'jumps' even for individuals from the same populations (Hedh & Hedenström 2016, Lislevand et al. 2017, Tomkovich et al. 2017. It is not known how individual migration schedules develop but it is plausible that juveniles from different natal sites mix in the wintering areas and then simply follow adults from their winter group to their breeding grounds. ...
Exploring the patterns of genetic structure in the context of geographical and phenotypic variation is important to understand the evolutionary processes involved in speciation. We investigated population and subspecies differentiation in the Common Ringed Plover Charadrius hiaticula, a high latitude wader that breeds in arctic and temperate zones from northeast Canada across Eurasia to the Russian Far East. Three subspecies, hiaticula, tundrae and psammodromus, are currently widely recognised, whereas a fourth subspecies, kolymensis, has been proposed based on geographic isolation and phenotypic differences. We genotyped 173 samples from eleven Common Ringed Plover breeding sites, representing all four putative subspecies, at eight polymorphic microsatellite loci to examine the patterns of population and subspecies differentiation. Bayesian clustering identified three genetic clusters among samples, corresponding to the breeding sites of the three currently recognised subspecies. The existence of the subspecies kolymensis was not supported. We also detected the presence of a previously unknown hybridisation zone extending from Northern Scandinavia to Belarus. Differentiation of the subspecies tundrae and hiaticula most likely occurred in allopatry on the Eurasian continent during past glaciation events, followed by population expansion leading to colonisation of Iceland and Greenland. The lack of genetic differentiation within the tundrae subspecies is consistent with ongoing range expansion and high gene flow maintained through migratory behaviour. We discuss the importance of historic climate changes, migratory behaviour and mating system on shaping the observed pattern of genetic differentiation.
... Tracking studies of two European populations have revealed that ringed plovers from southern Sweden utilize wintering sites along the whole European Atlantic coast, including Morocco (Hedh and Hedenström 2020), and ringed plovers from northern Norway winter along the coast of West Africa (Lislevand et al. 2017). Also, tracked populations breeding in Arctic Canada and sub-Arctic Russia have been shown to winter in West and East Africa, respectively (Tomkovich et al. 2017;Léandri-Breton et al. 2019). Apart from the differences in migration distances, temperate breeding populations generally experience longer breeding seasons, with frequent reports of successive clutches within monogamous pairs, and up to 2 months earlier egg-laying dates compared to populations breeding on more northerly latitudes, particularly those breeding in the Arctic (north of the Arctic Circle) and alpine areas within the species breeding range (Väisänen 1977;Pienkowski 1984a;Blomqvist et al. 2001;Wallander and Andersson 2003). ...
A common migratory pattern in birds is that northerly breeding populations migrate to more southerly non-breeding sites compared to southerly breeding populations ( leap-frog migration ). Not only do populations experience differences in migration distances, but also different environmental conditions, which may vary spatiotemporally within their annual cycles, creating distinctive selective pressures and migratory strategies. Information about such adaptations is important to understand migratory drivers and evolution of migration patterns. We use light-level geolocators and citizen science data on regional spring arrivals to compare two populations of common ringed plover Charadrius hiaticula breeding at different latitudes. We (1) describe and characterize the annual cycles and (2) test predictions regarding speed and timing of migration. The northern breeding population (NBP) wintered in Africa and the southern (SBP) mainly in Europe. The annual cycles were shifted temporally so that the NBP was always later in all stages. The SBP spent more than twice as long time in the breeding area, but there was no difference in winter. The NBP spent more time on migration in general. Spring migration speed was lower in the SBP compared to autumn speed of both populations, and there was no difference in autumn and spring speed in the NBP. We also found a larger variation in spring arrival times across years in the SBP. This suggests that a complex interaction of population specific timing and variation of breeding onset, length of breeding season, and proximity to the breeding area shape the annual cycle and migratory strategies.
Significance statement
Migration distance, climate, and the resulting composition of the annual cycle are expected to influence migration strategies and timing in birds. Testing theories regarding migration behaviours are challenging, and intraspecific comparisons over the full annual cycle are still rare. Here we compare the spatiotemporal distributions of two latitudinally separated populations of common ringed plovers using light-level geolocators. We found that there was a larger long-term variation in first arrival dates and that migration speed was slower only in spring in a temperate, short-distance migratory population, compared to an Arctic, long-distance migratory population. This suggests that a complex interaction of population specific timing and variation of breeding onset, length of breeding season and proximity to the breeding area shape the annual cycle and migratory behaviours.
... alpina), Wood Sandpiper (Tringa glareola), Graytailed Tattler (T. brevipes), Common Ringed Plover (Charadrius hiaticula), Amur Falcon (Falco amurensis), and Common Cuckoo (Cuculus canorus) (Zhang and Yang 1997, Meyburg et al. 2017, Tomkovich et al. 2017, David 2018. We speculate that there is a migration corridor between NE and E China and Myanmar and India for birds living in East Asia and the Indian subcontinent. ...
Little is known about the migration of the Gray-headed Lapwing (Vanellus cinereus). During the fall migration season of 2018, we tracked 3 birds from Guiyang Airport, China, a stopover site of Gray-headed Lapwing, in order to analyze their migratory and wintering movements by means of GPS-GSM satellite telemetry. We obtained 3 partial fall migration routes and 2 complete spring migration routes from September 2018 to July 2019. We identified E India and Bangladesh as their wintering ground and NE and E China as their breeding area. The spring migration covered 3,380 km and took 12 d on average (n = 2). Gray-headed Lapwings migrated at night, flying at speeds of 29.11–91.12 km/h. In winter, the mean home range area (95% utilization distribution) was 18.56 km2 (SD = 21.74, 3 birds) and the mean core area (50% utilization distribution) was 1.28 km2 (SD = 1.49, 3 birds).
... All rights reserved.' breeding site or other similar-sized species (Tomkovich et al. 2013, Lislevand and Hahn 2015, Brown et al. 2017, Johnson et al. 2017, Tomkovich et al. 2018. If so, it would be fascinating to understand the ecological and evolutionary drivers leading to such a spectrum of migratory strategies. ...
Effective conservation of migratory species depends on understanding both migratory connectivity and migration strategy. The Red‐necked Stint Calidris ruficollis is a small, highly migratory sandpiper of the East Asian‐Australasian Flyway, which is classified as Near Threatened due to ongoing population declines. We tracked the migration of three Red‐necked Stints breeding in southern Chukotka, Russia, using geolocators, and supplemented our tracking data with re‐sighting records of color‐flagged individuals. The three birds, all of which bred within 2km of each other, wintered in three different localities spanning nearly 5,000km. One individual completed its northward migration of >9400 km in two marathon flights; the second leg of that journey was completed in a nonstop flight of 5,350 km. The successful conservation of just this one population requires protection of wintering sites across a vast area, coupled with key staging sites along the flyway. We suggest that other migratory species may pose similar conservation challenges.
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... Our results are consistent with this general pattern and indicate that Canadian ringed plovers spend the winter at similar latitudes to the few northern Scandinavian breeding plovers tracked so far (Lislevand et al. 2016). Interestingly, individuals from the easternmost breeding population (Chukotka, Russia) and tracked with geolocators spent winter in the Arabian Peninsula and Northeast Africa (Tomkovich et al. 2017). This suggests an east-west divide in the species wintering distribution with western breeding birds wintering in West Africa and Europe and eastern breeding birds wintering in East Africa and Middle East. ...
Ecological barriers such as oceans, mountain ranges or glaciers can have a substantial influence on the evolution of animal migration. Along the migration flyway connecting breeding sites in the North American Arctic and wintering grounds in Europe or Africa, Nearctic species are confronted with significant barriers such as the Atlantic Ocean and the Greenland icecap. Using geolocation devices, we identified wintering areas used by Ringed Plovers nesting in the Canadian High‐Arctic and investigated migration strategies used by these Nearctic migrants along the transatlantic route. The main wintering area of the Ringed Plovers (n = 20) was located in Western Africa. We found contrasting seasonal migration patterns, with Ringed Plovers minimizing continuous flight distances over the ocean in spring by making a detour to stop in Iceland. In autumn, however, most individuals crossed the ocean in one direct flight from Southern Greenland to Western Europe, as far as Southern Spain. This likely resulted from prevailing anti‐clockwise winds associated with the Icelandic low‐pressure system. Moreover, the plovers we tracked largely circumvented the Greenland icecap in autumn, but in spring, some plovers apparently crossed the icecap above the 65°N. Our study highlighted the importance of Iceland as a stepping‐stone during the spring migration and showed that small Nearctic migrants can perform non‐stop transatlantic flights from Greenland to Southern Europe.
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Aim. The purpose of the paper is to analyze original data on rare birds in the poorly studied mountainous territory of the north of Transbaikalia. Methodology. The study was conducted mainly in the summer of 2019-2020 in mountain ranges: Udokan, Kodar, Kalarskiy and intermountain basins. Use was made of binoculars and photo registration. Results. Nine protected bird species (Red Data Books of Russia and Zabaikalsky krai) were observed: black-throated loon Gavia arctica, whooper swan Cygnus cygnus, harlequin duck Histrionicus histrionicus, crested honey buzzard Pernis ptilorhynchus, golden eagle Aquila chrysaetos, common crane Grus grus, Eurasian wren Troglodytes troglodytes, yellow-browed bunting Ocyris chrysophrys and yellow-breasted bunting O. aureoles. We also report the first confirmed case of nesting of lesser whitethroat Sylvia curruca and an unusual summer meeting of ringed plover Charadrius hiaticula in the Charskaya depression. For the lesser whitethroat, this is the easternmost point of the nesting area. Research implications. The data on the state of populations of rare species in mountainous areas in the north of Transbaikalia is important due to the threat of their extinction from the actively developing mining industry and the need to update the List of Protected Species of the Transbaikal Territory.
Discusses the geographical breeding origin and migration routes of 3.5 million waders using the Banc d'Arguin, Mauritania and Guinea-Bissau. For at least part of the ringed Charadrius hiaticula and grey plovers Pluvialis squatarola, redshanks Tringa totanus and to a lesser extent also bar-tailed godwit Limosa lapponica, whimbrels Numenius phaeopus, curlews Numenius arquatus and turnstones Arenaria interpres, there are indications that birds wintering in Guinea-Bissau originate from breeding areas further N and E, as compared to birds wintering in Mauritania. During spring migration most wader species wintering in W African coastal wetlands travel via a route following the W African and W European coastline. Little stint Calidris minuta, curlew sandpiper Calidris ferruginea and curlew are exceptions to this rule. There are indications that at least part of these birds migrate through NW Africa, the Mediterranean and the Black Sea area. -Authors
Semipalmated Sandpiper (Calidris pusilla) populations have undergone significant declines at core nonbreeding sites in northeastern South America. Breeding populations have also declined in the eastern North American Arctic, but appear to be stable or increasing in the central and western Arctic. To identify vulnerable populations and sites, we documented the migratory connectivity of Semipalmated Sandpipers using light-level geolocators, deploying 250 at 8 Arctic sites across the species' breeding range from 2011 to 2015, plus 87 at a single wintering site in northeastern Brazil in 2013 and 2014. We recovered 59 units and resighted 7 more (26% return rate) on the breeding grounds, but none at the nonbreeding site. We recovered only ∼3% of units deployed in 2013 at eastern Arctic breeding sites, but recovered 33% of those deployed in 2015. Overall, birds with geolocators were 57% as likely to return as those carrying alphanumeric flags. Stopover durations at prairie sites (mean: 8.7 days southbound, 6.7 days northbound) were comparable with durations estimated by local banding studies, but geolocator-tagged birds had longer stopovers than previously estimated at James and Hudson Bay, the Bay of Fundy, and the Gulf of Mexico. Migration routes confirmed an eastern Arctic connection with northeastern South America. Birds from eastern Alaska, USA, and far western Canada wintered from Venezuela to French Guiana. Central Alaskan breeders wintered across a wider range from Ecuador to French Guiana. Birds that bred in western Alaska wintered mainly on the west coasts of Central America and northwestern South America, outside the nonbreeding region in which population declines have been observed. Birds that bred in the eastern Arctic and used the Atlantic Flyway wintered in the areas in South America where declines have been reported, whereas central Arctic-breeding populations were apparently stable. This suggests that declines may be occurring on the Atlantic Flyway and in the eastern Arctic region.
Costs of migration, in terms of time, energy, and mortality risk, have a strong theoretical and empirical foundation in the study of birds. We expect these costs to be most severe for extreme long-distance migratory landbirds, whose demanding annual routines (e.g. non-stop flights 8000 km and return journeys 30 000 km) may approach their maximum physiological capabilities. To explore whether this is true, we review evidence in long-jump migratory shorebirds (Scolopacidae), focusing most on the prototypical example, the Alaska-breeding bar-tailed godwit Limosa lapponica baueri. Contrary to expectations, these and similar birds demonstrate high adult survival, little evidence for elevated mortality during migration, no apparent minimisation of non-stop flight distances, and low inter-and intra-individual variation in migration performance. Two key aspects of extreme migrants may explain these findings: 1) a counter-intuitively conservative annual-cycle strategy, which minimises risks and enables dissipation of carry-over effects before fitness consequences arise; and 2) selection pressure during early life, which quickly removes low-performing individuals from the population. We hypothesise that these two factors, applicable to extreme strategies in a wide range of taxa, act to truncate the range of individual quality in a population, and decrease the prevalence and detectability of carry-over effects. Testing these hypotheses is challenging, as it requires comparative studies of demography and individual quality spectra along a continuum of extremeness. However, it has important potential implications for interpreting individual variation, designing studies of cross-seasonal interactions or costs of migration, and recognising early-warning signs of population decline. For example, the most extreme shorebird migrations rely on abundant but difficult-to-access resources; the high minimum individual performance required for survival predicts that degradation of these resource hot-spots will propel rapid population collapse, rather than incremental declines in condition or performance. Therefore, in extreme migrants, we may paradoxically view populations as operating close to the edge, even while individuals are not.
The Great Knot Calidris tenuirostris is one of the iconic long-distance migratory species of the East Asian-Australasian Flyway. However, despite extensive flagging and banding efforts, very little is known about the migratory strategies and the breeding grounds of this species that spends the non-breeding season mainly on the northern shorelines of Australia. Using light-level geolocators deployed on Great Knots at Roebuck Bay (Western Australia), we describe the individual migration strategies, breeding locations and breeding-related behaviour. Based on data from eight successfully tracked individuals, we found that all except one migrated to the western part of the known breeding range. This was 2,000–2,500 km from the eighth individual that commenced breeding in the potentially separated eastern part of the range. Light intensity and temperature profiles provided evidence that four of the birds successfully hatched chicks. Of the three which failed, one appeared to have laid a second clutch before failing again. Arrival at the breeding grounds and the laying of eggs were remarkably synchronous between individuals, as were the arrival dates back at Roebuck Bay. Departure from the breeding grounds was more spread out, partly dependent on breeding success and also as a result of females probably leaving the nesting area before males. The individual migration strategies confirmed the strong dependence of this species on the Yellow Sea as their major stopover site during both southward and northward migration. Furthermore, all individuals stopped at least once on their northward journey to the Yellow Sea from Australia. And in reverse, all individuals stopped at least once on the southward migration before arriving at the Yellow Sea coming from their Arctic breeding grounds. The results indicate that this species will most likely be further affected by the rapid habitat loss in the area of the Yellow Sea and other parts of the Chinese coastline.
Landbirds undertaking within-continent migrations have the possibility to stop en route, but most long-distance migrants must also undertake large non-stop sea crossings, the length of which can vary greatly. For shorebirds migrating from Iceland to West Africa, the shortest route would involve one of the longest continuous sea crossings while alternative, mostly overland, routes are available. Using geolocators to track the migration of Icelandic whimbrels (Numenius phaeopus), we show that they can complete a round-trip of 11,000 km making two non-stop sea crossings and flying at speeds of up to 24 m s−1; the fastest recorded for shorebirds flying over the ocean. Although wind support could reduce flight energetic costs, whimbrels faced headwinds up to twice their ground speed, indicating that unfavourable and potentially fatal weather conditions are not uncommon. Such apparently high risk migrations might be more common than previously thought, with potential fitness gains outweighing the costs.
How individual birds schedule their movements and use different sites during the non-breeding season are fundamental issues in avian migration ecology, and studies have often revealed strong seasonal variation in such strategies. Using geolocators we tracked Common Ringed Plovers Charadrius hiaticula from northern Norway to West Africa and back to assess whether there were differences in migratory speed, duration and stopover use between autumn and spring migration and whether birds used multiple sites during the non-breeding season. Although the pace of migration was similar between autumn and spring, the length of flight bouts and duration of the preceding stopovers were positively correlated only in autumn. Four of five birds showed a marked southward movement in mid-winter. This article is protected by copyright. All rights reserved.
Many shorebird populations have declined, and mapping habitat use during the
non-breeding period and determining timing of use have become increasingly
urgent for conservation. This study provides the first detailed location information
on the complete migration cycle for the subspecies of Dunlin Calidris alpina
sakhalina breeding in Chukotka, Russia. Geolocators were deployed and recovered
on breeding Dunlin at Chaun River Delta (n = 12 of 35 deployed) and Belyaka
Spit, Russia (n = 11 of 25 deployed), between 2011 and 2014. Dunlin migrated
south through the Sea of Okhotsk to the Korean Peninsula and China. Upon
reaching the Yellow Sea, ten individuals stopped for 39–74 (mean ± SE; 54 ± 4)
days before moving on to wintering grounds in China, and two remained in the
area throughout the winter. Tagged Dunlin wintered in China, South Korea, and
Vietnam, primarily between the northwest Yellow Sea and the Gulf of Tonkin in
China. Nine individuals moved from their wintering grounds north to areas
around the Yellow Sea between 10 March and 3 April (mean: 23 March), where
they spent 23–73 (53 ± 4) days before continuing to the Arctic breeding grounds
with only short stopovers (<10 days). Dunlin may be site-faithful to their winter
and stopover areas; geolocators on three individuals recorded >1 year of data
and indicated very similar migratory routes and use of winter areas between
years. At the scale of this study, there was no segregation of Dunlin that bred in
the Chaun River Delta and at Belyaka Spit during the non-breeding season.
Dunlin breeding in Chukotka, Russia, are particularly reliant on coastal habitat
around the Yellow Sea, as well as the Fujian coast and Yangtze River floodplain in
China for wintering habitat. The Yellow Sea is an important stopover site during
both southern and northern migrations, as are coastal areas around northern
Sakhalin Island and the Sea of Okhotsk.
Background
Geolocators are useful for tracking movements of long-distance migrants, but potential negative effects on birds have not been well studied. We tested for effects of geolocators (0.8–2.0 g total, representing 0.1–3.9 % of mean body mass) on 16 species of migratory shorebirds, including five species with 2–4 subspecies each for a total of 23 study taxa. Study species spanned a range of body sizes (26–1091 g) and eight genera, and were tagged at 23 breeding and eight nonbreeding sites. We compared breeding performance and return rates of birds with geolocators to control groups while controlling for potential confounding variables.
Results
We detected negative effects of tags for three small-bodied species. Geolocators reduced annual return rates for two of 23 taxa: by 63 % for semipalmated sandpipers and by 43 % for the arcticola subspecies of dunlin. High resighting effort for geolocator birds could have masked additional negative effects. Geolocators were more likely to negatively affect return rates if the total mass of geolocators and color markers was 2.5–5.8 % of body mass than if tags were 0.3–2.3 % of body mass. Carrying a geolocator reduced nest success by 42 % for semipalmated sandpipers and tripled the probability of partial clutch failure in semipalmated and western sandpipers. Geolocators mounted perpendicular to the leg on a flag had stronger negative effects on nest success than geolocators mounted parallel to the leg on a band. However, parallel-band geolocators were more likely to reduce return rates and cause injuries to the leg. No effects of geolocators were found on breeding movements or changes in body mass. Among-site variation in geolocator effect size was high, suggesting that local factors were important.
Conclusions
Negative effects of geolocators occurred only for three of the smallest species in our dataset, but were substantial when present. Future studies could mitigate impacts of tags by reducing protruding parts and minimizing use of additional markers. Investigators could maximize recovery of tags by strategically deploying geolocators on males, previously marked individuals, and successful breeders, though targeting subsets of a population could bias the resulting migratory movement data in some species.
Lake Baikal lies in eastern Siberia, Russia. Due to its huge size, its waterbird fauna is still insufficiently known in spite of a long history of relevant research and the efforts of local and visiting ornithologists and birdwatchers. Overall, 137 waterbird species have been recorded at Lake Baikal since 1800, with records of five further species considered not acceptable, and one species recorded only prior to 1800. Only 50 species currently breed at Lake Baikal, while another 11 species bred there in the past or were recorded as occasional breeders. Only three species of conservation importance (all Near Threatened) currently breed or regularly migrate at Lake Baikal : Asian Dowitcher Limnodromus semipalmatus, Black-tailed Godwit Limosa limosa and Eurasian Curlew Numenius arquata.
The Common Ringed Plover Charadrius hiaticula has a widespread breeding distribution at temperate and high latitudes in Europe and NE Canada. Three subspecies have been identified: C. h. psammodroma has a breeding range from the Faeroes to NE Canada, C. h. hiaticula breeds in Britain, S Scandinavia, Northern and Eastern Europe and C. h. tundrae breeds from N Scandinavia to Russia. These populations have a wintering range from Britain to S Africa and SW Asia, where psammodroma and tundrae leap-frog most of the hiaticula race by breeding further north and wintering further south. Even though the non-breeding distribution of the Ringed Plover is quite well known, knowledge of movements of the subspecies from breeding grounds to wintering sites is sparse. The Icelandic population (psammodroma) has been thought to winter in S Europe and NW Africa but only one winter ringing recovery has been reported, in S Spain. Here we present data on 104 recoveries of Icelandic Ringed Plovers outside Iceland, 86 of colour-ringed birds and 18 of metal-ringed birds. Most recoveries in autumn and spring were from Britain, Ireland and France. In winter most recoveries were from S Europe and NW Africa. Icelandic Ringed Plovers leap-frog the British population and probably a part of the S Scandinavia population.
Based on ring recoveries and observations of colour-marked individuals, we identify the non-breeding range, migration routes and staging areas used by Red Knots Calidris canutus of the subspecies rogersi from its southernmost sub-arctic breeding population in Chukotka, NE Asia. We also present the first three geolocator tracks of individual rogersi adult males. The non-breeding grounds of rogersi extend from New Zealand to NW Australia, with the majority going to the eastern part of this range (primarily New Zealand and E Australia). Three key staging areas for migrants are indicated by the geolocator data: (1) the NW Yellow Sea, (2) the western Sea of Okhotsk, north of the Amur River delta, and (3) the Gulf of Carpentaria, Australia. The Yellow Sea is extensively used during both north and south migration, whereas the other two sites are only used during south migration. The three birds with geolocators made round trips of 23,000–31,000 km. The two that went to New Zealand made direct flights of about 10,100 km from New Zealand to the NW Yellow Sea, the longest non-stop flight known for Red Knots to date. Mostly the flight paths taken by the three birds between stopovers were not further from the great circle route that the ca. 200 km margin of error associated with geolocator fixes. The only exception was that in most cases they appeared to fly south of the Kamchatka Peninsula when migrating between their breeding grounds and the Yellow Sea (four of five flight paths passed south around Kamchatka).
We review the use of light-level geolocators to track long-distance migrant shorebirds and describe several techniques that an analyst can employ to obtain the best possible estimates of a bird’s location. These include: calibration using perfectly bright and clear days, longitude averaging, point clusters, habitat knowledge, the use of resightings of the birds, dealing with the different error patterns before and after the spring and fall equinoxes, weather pattern correction, the effects of artificial light, interpretation of geolocator fixes obtained during migratory flights, the use of data recorded by a geolocator’s conductivity sensor to determine flight duration and location estimation when a bird is in the Arctic using the nightshade technique. We also review the accuracy that can be expected in shorebird studies using geolocators.
Waterfowl (Anseriformes) and shorebirds (Charadriiformes) are the most common wild vectors of influenza A viruses. Due to their migratory behavior, some may transmit disease over long distances. Migratory connectivity studies can link breeding and nonbreeding grounds while illustrating potential interactions among populations that may spread diseases. We investigated Dunlin (Calidris alpina), a shorebird with a subspecies (C. a. arcticola) that migrates from nonbreeding areas endemic to avian influenza in eastern Asia to breeding grounds in northern Alaska. Using microsatellites and mitochondrial DNA, we illustrate genetic structure among six subspecies: C. a. arcticola, C. a. pacifica, C. a. hudsonia, C. a. sakhalina, C. a. kistchinski, and C. a. actites. We demonstrate that mitochondrial DNA can help distinguish C. a. arcticola on the Asian nonbreeding grounds with >70% accuracy depending on their relative abundance, indicating that genetics can help determine if C. a. arcticola occurs where they may be exposed to highly pathogenic avian influenza (HPAI) during outbreaks. Our data reveal asymmetric intercontinental gene flow, with some C. a. arcticola short-stopping migration to breed with C. a. pacifica in western Alaska. Because C. a. pacifica migrates along the Pacific Coast of North America, interactions between these subspecies and other taxa provides route for transmission of HPAI into other parts of North America.This article is protected by copyright. All rights reserved.
A major aim of bird ringing is to provide information about the migration and movements of bird populations. However, in comparison with demographic studies, little research has been devoted to improving quantitative inferences through large‐scale spatial analyses. This represents a serious knowledge gap because robust information on geographical linkages of migratory populations throughout the annual cycle is necessary to understand the ecology and evolution of migrants and for the conservation and management of populations.
Here, we review recent developments and emerging opportunities for the quantitative study of movements of bird populations based on marked birds. Large‐scale spatial analyses of ringing data need to account for spatiotemporal variation in re‐encounter probability and the complexity of movement processes, including variability among individuals and populations in migration direction and distance.
We identify seven recent studies that used quantitative methods for large‐scale spatial analyses of ringing and re‐encounter data gathered by national ringing centres. In most cases, numbers ringed and recovered in a series of source and destination areas were used to derive estimates of the proportion of each source population travelling to each destination area. Where recovery data were sparse, precision was improved by incorporating information on re‐encounter probabilities of similar species. When numbers ringed were not available, inferences could sometimes be drawn based on local recapture data from the source areas.
Studies to date illustrate that analyses of these large‐scale ringing data sets can provide robust quantitative inferences. Further work is needed to develop these modelling approaches and to test their sensitivity to key assumptions using both real and simulated data. Data for all birds that were marked, not only those re‐encountered, are often inaccessible and should be computerised in parallel with analytical developments. Further, there is great potential for the formal combination of re‐encounter data with information from additional data sources such as counts and detailed movement data from radiotracking or data loggers. Because data from bird ringing operations cover long periods of time and exist in large quantities, they hold great promise for inferring spatiotemporal migration patterns, including changes in relation to climate, land use change and other environmental drivers.
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.
Because museum scientists and conservationists are natural allies in the struggle to preserve biodiversity, conflict over the legality, morality, and value of collecting scientific specimens is counterproductive. Modern bird specimens contain a variety of data, summarized briefly herein, that are applied to numerous questions concerning the biology of birds, many of which have direct and often critical relevance to conservation. In particular, continued collecting of specimens has been shown to be critical in determining species-level classification in birds; unless species limits are established correctly, conservation priorities cannot be established reliably. Objections to collecting specimens are summarized and discussed. Calculations are presented to show that the effect of collecting specimens on most bird populations is insignificant. Moral objections t o collecting specimens seem to reflect a lack of awareness of the extent and causes of natural mortality, as well as a failure to recognize the magnitude of unintentional mortality inflicted on bird populations by routine human activities. The reason why more specimens are needed than currently exist in museum collections is that most existing specimens lack the data needed for most kinds of modern analyses, and even common species are represented by inadequate samples for research. Reasons are given for why equivalent data cannot be obtained solely from living birds that are subsequently released. Objecting to collecting specimens because it sets a bad example for developing countries trying to establish an environmental ethic is counterproductive in that it draws attention away from the fundamental units of concern for conservation biology: the population, and the habitat that supports it. Biological specimens differ from some other scientific specimens (e.g. archaeological) in that they are renewable resources whose removal does not deplete a country's national heritage. Misconceptions about museum scientists and their motives are discussed. Regarding collecting permits, recommendations are presented concerning (1) numbers of specimens, (2) percentage of specimens left in the host country, (3) species composition, (4) deposition of specimens, and (5) processing permit applications. Regulating agencies are often overly enthusiastic i n restricting scientific collecting, which is the only kind of mortality that is so highly controlled and yet from which bird species might derive benefit, whereas the same or sister agencies often permit and even encourage activities that are responsible for massive mortality in bird populations. Given that (1) the goal of scientists, conservation agencies, and governments is protection of populations, not individual birds; (2) scientific collecting has no measurable impact on the vast majority of bird populations; (3) scientific specimens represent an important source of information on bird biology and conservation; and (4) existing scientific collections are largely inadequate for answering many questions that could be answered with greater numerical, seasonal, or geographic representation, then it follows that continued scientific collecting will benefit ornithology and conservation and should, therefore, be encouraged by conservation and government agencies.
In this paper we review information on the status, population size and migrations of waders within southern and eastern Africa, western Asia and the USSR, in an attempt to show where major concentrations occur, and the routes taken to and from their Palearctic breeding grounds. For much of this area, the information is meagre. The best-surveyed areas are Iran, the Nile Delta in Egypt, Kenya, and the coasts of Turkey, Namibia and South Africa. Rough estimates are available also for Sudan and the Gulf coast of Saudi Arabia. Large numbers of wintering waders occur in the Persian Gulf, Nile Delta, White Nile in Sudan, Lake Chad and the Rift Valley lakes of Ethiopia and Kenya. Long-term changes in the seasonal rains of Africa make it difficult to make representative population estimates. Coastal surveys in Kenya, Tanzania, South Africa and Namibia have shown that large numbers of certain species occur on rocky and coral shores, sandy beaches and coastal inlets. Observations and ringing recoveries indicate that there are several routes taken by migrating waders through Africa: along the west coast of southern Africa to the Gulf of Guinea and then across the Sahara to the Mediterranean; along the Rift Valley lakes and the River Nile; and along the east coast of Africa. In view of the large numbers of wintering waders in the Nile Delta and Persian Gulf it is likely that these areas are important stop-over points for migrants, whilst ringing recoveries in the Black Sea and Caspian Sea show that these areas are also used. By highlighting the huge gaps in the knowledge of waders in these parts of the world, we hope to stimulate and direct waderologists to the poorer-known parts of Africa and Asia.
Abstract It has been suggested that birds migrate faster in spring than in autumn because of competition for arrival order at breeding grounds and environmental factors such as increased daylight. Investigating spring and autumn migration performances is important for understanding ecological and evolutionary constraints in the timing and speed of migration. We compiled measurements from tracking studies and found a consistent predominance of cases showing higher speeds and shorter durations during spring compared to autumn, in terms of flight speeds (airspeed, ground speed, daily travel speed), stopover duration, and total speed and duration of migration. Seasonal differences in flight speeds were generally smaller than those in stopover durations and total speed/duration of migration, indicating that rates of foraging and fuel deposition were more important than flight speed in accounting for differences in overall migration performance. Still, the seasonal differences in flight speeds provide important support for time selection in spring migration.
Basic questions about the life histories of migratory birds have confounded scientists for generations, yet we are nearing an era of historic discovery as new tracking technologies make it possible to determine the timing and routes of an increasing number of bird migrations. Tracking small flying animals as they travel over continental-scale distances is a difficult logistical and engineering challenge. Although no tracking system works well with all species, improvements to traditional technologies, such as satellite tracking, along with innovations related to global positioning systems, cellular networks, solar geolocation, radar, and information technology are improving our understanding of when and where birds go during their annual cycles and informing numerous scientific disciplines, including evolutionary biology, population ecology, and global change. The recent developments described in this article will help us answer many long-standing questions about animal behavior and life histories.
Places where migrant birds stop to rest, drink, and eat at are often described as either stopover or staging sites. Attempts have been made to differentiate between these two terms but they are frequently used interchangeably. Some authors have equated staging sites with sites that attract large concentrations (many thousands) of birds, a definition that others have expanded to include long stopover durations and significant rates of refueling on predictable, abundant prey. It has also been suggested that birds using staging sites are those that employ a jumping strategy during migration. I argue that while all sites where birds rest and feed during migration are stopover sites, further classification of stopover sites is of ecological and conservation value. I propose that sites with abundant, predictable food resources where birds prepare for an energetic challenge (usually a long flight over a barrier such as an ocean or a desert) requiring substantial fuel stores and physiological changes without which significant fitness costs are incurred are most appropriately described as staging sites.
1. Geolocation by light allows for tracking animal movements, based on measurements of light intensity over time by a data‐logging device (‘geolocator’). Recent developments of ultra‐light devices (<2 g) broadened the range of target species and boosted the number of studies using geolocators. However, an inherent problem of geolocators is that any factor or process that changes the natural light intensity pattern also affects the positions calculated from these light patterns. Although the most important factors have been identified, estimation of their effect on the accuracy and precision of positions estimated has been lacking but is very important for the analyses and interpretation of geolocator data.
2. The ‘threshold method’ is mainly used to derive positions by defining sunrise and sunset times from the light intensity pattern for each recorded day. This method requires calibration: a predefined sun elevation angle for estimating latitude by fitting the recorded day/night lengths to theoretical values across latitudes. Therewith, almost constant shading can be corrected for by finding the appropriate sun elevation angle.
3. Weather, topography and vegetation are the most important factors that influence light intensities. We demonstrated their effect on the measurement of day/night length, time of solar midnight/noon and the resulting position estimates using light measurements from stationary geolocators at known places and from geolocators mounted on birds. Furthermore, we investigated the influence of different calibration methods on the accuracy of the latitudinal positions.
4. All three environmental factors can influence the light intensity pattern significantly. Weather and an animal’s behaviour result in increased noise in positioning, whereas topography and vegetation result in systematic shading and biased positions. Calibration can significantly shift the estimated latitudes and potentially increase the accuracy, but detailed knowledge about the particular confounding factors and the behaviour of the studied animal is crucial for the choice of the most appropriate calibration method.
Long-distance migration has evolved in many organisms moving through different media and using various modes of locomotion and transport. Migration continues to evolve or become suppressed as shown by ongoing dynamic and rapid changes of migration patterns. This great evolutionary flexibility may seem surprising for such a complex attribute as migration. Even if migration in most cases has evolved basically as a strategy to maximise fitness in a seasonal environment, its occurrence and extent depend on a multitude of factors. We give a brief overview of different factors (e.g. physical, geographical, historical, ecological) likely to facilitate and/or constrain the evolution of long-distance migration and discuss how they are likely to affect migration. The basic driving forces for migration are ecological and biogeographic factors like seasonality, spatiotemporal distributions of resources, habitats, predation and competition. The benefit of increased resource availability will be balanced by costs associated with the migratory process in terms of time (incl. losses of prior occupancy advantages), energy and mortality (incl. increased exposure to parasites). Furthermore, migration requires genetic instructions (allowing substantial room for learning in some of the traits) about timing, duration and distance of migration as well as about behavioural and physiological adaptations (fuelling, organ flexibility, locomotion, use of environmental transport etc) and control of orientation and navigation. To what degree these costs and requirements put constraints on migration often depends on body size according to different scaling relationships. From this exposé it is clear that research on migration warrants a multitude of techniques and approaches for a complete as possible understanding of a very complex evolutionary syndrome. In addition, we also present examples of migratory distances in a variety of taxons. In recent years new techniques, especially satellite radio telemetry, provide new information of unprecedented accuracy about journeys of individual animals, allowing re-evaluation of migration, locomotion and navigation theories.
Shorebirds, or waders, form an ecologically (but not phylogenetically) homogenous group of birds that, despite this homogeneity,
exhibits clear correlated contrasts in habitat use and migration distance between closely related species pairs. In addition,
within species there is distinct variation in breeding and wintering latitudes, i.e. migration distance. I examine here such
contrasts at different taxonomic levels and evaluate what we can learn about selective forces on habitat selection and the
evolution of migration strategies in birds. My primary example is the worldwide migration system of the Red Knot Calidris canutus. These sandpipers breed only on high arctic tundra (65–83°N), but they move south from their disjunct, circumpolar breeding
areas to nonbreeding sites on the coasts of all continents (except Antarctica), between latitudes 58°N and 53°S. Due to their
specialized sensory capabilities, Red Knots generally eat hard-shelled prey found on intertidal, mostly soft, substrates.
As a consequence, ecologically suitable coastal sites are few and far between, so they must routinely undertake flights of
many thousands of kilometres. In contrast to prediction, Red Knots at tropical intertidal sites have lower fuelling rates
than birds at more southern or northern latitudes. This leads to greater time–stress in the southernmost wintering populations
that not only have to cover over 14,000km in single migrations, but also cannot rely on tropical regions to make refuelling
stops. Rapid human-caused losses of the food-base in staging areas during both north- and southward migrations have been demonstrated
to have caused rapid declines in several Red Knot populations. Detailed worldwide eco-demographic research on these extreme
long-distance migrants, as embodied in, for example, the recently established Global Flyway Network, yields a two-pronged
benefit: (1) on the basis of the unintended large-scale experiments carried out by humans, we rapidly come to grips with the
selection pressures moulding the migration strategies of migrant birds, and (2) in applied contexts, the work gives instantaneous
feedbacks on the conservation consequences of man-made alterations to wetland environments at the relevant global spatial
scales.
Migration is the regular seasonal movement of animals from one place to another, often from a breeding site to a nonbreeding site and back. Because the act of migration makes it difficult to follow individuals and populations year round, our understanding of the ecology and evolution of migrating organisms, particularly birds, has been severely impeded. Exciting new advances in satellite telemetry, genetic analyses and stable isotope chemistry are now making it possible to determine the population and geographical origin of individual birds. Here, we review these new approaches and consider the relevance of understanding migratory connectivity to ecological, evolutionary and conservation issues.
The northern wheatear (Oenanthe oenanthe) is a small (approx. 25 g), insectivorous migrant with one of the largest ranges of any songbird in the world, breeding from the eastern Canadian Arctic across Greenland, Eurasia and into Alaska (AK). However, there is no evidence that breeding populations in the New World have established overwintering sites in the Western Hemisphere. Using light-level geolocators, we demonstrate that individuals from these New World regions overwinter in northern sub-Sahara Africa, with Alaskan birds travelling approximately 14 500 km each way and an eastern Canadian Arctic bird crossing a wide stretch of the North Atlantic (approx. 3500 km). These remarkable journeys, particularly for a bird of this size, last between one to three months depending on breeding location and season (autumn/spring) and result in mean overall migration speeds of up to 290 km d(-1). Stable-hydrogen isotope analysis of winter-grown feathers sampled from breeding birds generally support the notion that Alaskan birds overwinter primarily in eastern Africa and eastern Canadian Arctic birds overwinter mainly in western Africa. Our results provide the first evidence of a migratory songbird capable of linking African ecosystems of the Old World with Arctic regions of the New World.
Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64° N in Bering Strait up to latitude 75° N far north of Wrangel Island, with eastward flights making up 79% of all track directions.
The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian–American bird migration system in a wide Arctic sector between longitudes 110° E and 130° W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3–5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40–80 degrees of longitude, corresponding to distances and durations of 1400–2600 km and 26–48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.
This book presents an up-to-date, detailed and thorough review of the most fascinating ecological findings of bird migration. It deals with all aspects of this absorbing subject, including the problems of navigation and vagrancy, the timing and physiological control of migration, the factors that limit their populations, and more. Author, Ian Newton, reveals the extraordinary adaptability of birds to the variable and changing conditions across the globe, including current climate change. This adventurous book places emphasis on ecological aspects, which have received only scant attention in previous publications. Overall, the book provides the most thorough and in-depth appraisal of current information available, with abundant tables, maps and diagrams, and many new insights. Written in a clear and readable style, this book appeals not only to migration researchers in the field and Ornithologists, but to anyone with an interest in this fascinating subject. * Hot ecological aspects include: various types of bird movements, including dispersal and nomadism, and how they relate to food supplies and other external conditions * Contains numerous tables, maps and diagrams, a glossary, and a bibliography of more than 2,700 references * Written by an active researcher with a distinguished career in avian ecology, including migration research.
1. Migrants that leave the Palaearctic Region in winter are in the main the most insectivorous species. In Britain nearly one-third of the breeding passerine birds depend for winter-quarters on Africa south of the Sahara. 2. The Western Palaearctic has the potential wintering ground provided by tropical Africa, twice its own size, due south of it. On the other hand, the potential wintering grounds south of the Central and the Far Eastern Palaearctic are far smaller than these are. Consequently much of Palaearctic migration is directed south-west in autumn; Africa is sought by nearly as large a proportion of birds breeding in the Central as in the Western Palaearctic and even by some Far Eastern birds. 3. Some of the migrant species reaching Africa winter only north of the equator, some south, some throughout. Ecological reasons for these winter ranges are rarely obvious. 4. In most species populations breeding in different areas are not segregated in winter. 5. In the area between the Sahara and the equator the migrants arrive after the annual rains and experience increasing desiccation throughout their stay. Although few of the resident birds are breeding at that season it is surprising that so many immigrants can be supported under these severe conditions. 6. South of the equator the migrants arrive in time for the annual flush of vegetation, when the local birds are beginning to breed; so that both resident and immigrant populations reach their peak together--at the time of apparent maximum of food. 7. In some localities and in some genera the immigrant birds outnumber the residents. 8. Details of ecological adjustment in winter-quarters need to be worked out. Many immigrant birds appear to share a (possibly superabundant) food supply with allied resident species; some are highly itinerant. At least one species finds an unoccupied niche. 9. Africa has probably been an important wintering area since early in the Tertiary. The numbers involved varied most during the Pleistocene, with only one-third as many birds at a glacial maximum as during a climatic optimum. The post-glacial changes in the migrant flood have also been great, owing to the effects of human activity.
Ringing recoveries, simulated recoveries and observational data have been used to describe the migration of the Ringed Plover. The analysis indicates the presence of eight wintering regions. These are, the British Isles; southern France and northern Spain; the Iberian peninsula; north west Africa; north east Africa; the Arabian peninsula; western and southern Africa; south east Asia. Allopatric breeding populations occupy the same regions in winter (synheimic) throughout much of the northern and western portions of the winter range. The phenomenon of "leap frog" migration exists, but is less distinct than reported by earlier studies.
Methods for determining patterns of migratory connectivity in animal ecology have historically been limited due to logistical challenges. Recent progress in studying migratory bird connectivity has been made using genetic and stable-isotope markers to assign migratory individuals to their breeding grounds. Here, we present a novel Bayesian approach to jointly leverage genetic and isotopic markers and we test its utility on two migratory passerine bird species. Our approach represents a principled model-based combination of genetic and isotope data from samples collected on the breeding grounds and is able to achieve levels of assignment accuracy that exceed those of either method alone. When applied at large scale the method can reveal specific migratory connectivity patterns. In Wilson's warblers (Wilsonia pusilla), we detect a subgroup of birds wintering in Baja that uniquely migrate preferentially from the coastal Pacific Northwest. Our approach is implemented in a way that is easily extended to accommodate additional sources of information (e.g. bi-allelic markers, species distribution models, etc.) or adapted to other species or assignment problems.
The numerical and functional significance of the Wadden Sea over the past 30 years for twelve wader species is analysed. These species are: Ringed Plover Charadrius hiaticula, Grey Plover Pluvialis squatarola, Red Knot Calidris canutus, Sanderling Calidris alba, Curlew Sandpiper Calidris ferruginea, Dunlin Calidris alpina, Bar-tailed Godwit Limosa lapponica, Eurasian Curlew Numenius arquata, Spotted Redshank Tringa erythropus, Redshank Tringa totanus, Greenshank Tringa nebularia, and Ruddy Turnstone Arenaria interpres. The results of wader counts and wader catching activities are described per species. For every species attention is paid to the spatial distribution within the area, the seasonal changes, and the changes over the past 30 years with respect to the numbers of waders. The population composition is analysed with respect to the geographical breeding origin as deduced from morphometrics, sex ratio and/or age composition, and the changes in primary moult and body mass in the course of the non-breeding season. Findings with respect to the breeding origins were obtained by comparing the wing- and culmen lengths of waders captured in the Wadden Sea with the measurements of skins collected in a series of zoological museums at the northern hemisphere.
Bird migration and orientation at high latitudes are of special interest because of the difficulties associated with different compass systems in polar areas and because of the considerable differences between flight routes conforming to loxodromes (rhumblines) or orthodromes (great circle routes). Regular and widespread east-north-east migration of birds from the northern tundra of Siberia towards North America across the Arctic Ocean (without landmark influences) were recorded by ship-based tracking radar studies in July and August. Field observations indicated that waders, including species such as Phalaropusfulicarius and Calidris melanotos, dominated, but also terns and skuas may have been involved. Analysis of flight directions in relation to the wind showed that these movements are not caused by wind drift. Assuming possible orientation principles based on celestial or geomagnetic cues, different flight trajectories across the Arctic Ocean were calculated: geographical loxodromes, sun compass routes, magnetic loxodromes and magnetoclinic routes. The probabilities of these four alternatives are evaluated on the basis of both the availability of required orientation cues and the predicted flight paths. This evaluation supports orientation along sun compass routes. Because of the longitudinal time displacement sun compass routes show gradually changing compass courses in close agreement with orthodromes. It is suggested that an important migration link between Siberia and North American stopover sites 1000-2500km apart across the Arctic Ocean has evolved based on sun compass orientation along orthodrome-like routes.
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