Individuality in bird migration: routes and timing

The exploration of animal migration has entered a new era with individual-based tracking during multiple years. Here, we investigated repeated migratory journeys of a long-distance migrating bird, the marsh harrier Circus aeruginosus, in order to analyse the variation within and between individuals with respect to routes and timing. We found that there was a stronger individual repeatability in time than in space. Thus, the annual timing of migration varied much less between repeated journeys of the same individual than between different individuals, while there was considerable variation in the routes of the same individual on repeated journeys. The overall contrast in repeatability between time and space was unexpected and may be owing to strong endogenous control of timing, while short-term variation in environmental conditions (weather and habitat) might promote route flexibility. The individual variation in migration routes indicates that the birds navigate mainly by other means than detailed route recapitulation based on landmark recognition.     

The strength of migratory connectivity for birds en route to breeding through the Gulf of Mexico

The strength of migratory connectivity is a measure of the cohesion of populations among phases of the annual cycle, including breeding, migration, and wintering. Many Nearctic‐Neotropical species have strong migratory connectivity between breeding and wintering phases of the annual cycle. It is less clear if this strength persists during migration when multiple endogenous and exogenous factors may decrease the cohesion of populations among routes or through time along the same routes. We sampled three bird species, American redstart Setophaga ruticilla, ovenbird Seiurus aurocapilla, and wood thrush Hylocichla mustelina, during spring migration through the Gulf of Mexico region to test if breeding populations differentiate spatially among migration routes or temporally along the same migration routes and the extent to which within‐population timing is a function of sex, age, and carry‐over from winter habitat, as measured by stable carbon isotope values in claws (δ¹³C). To make quantitative comparisons of migratory connectivity possible, we developed and used new methodology to estimate the strength of migratory connectivity (MC) from probabilistic origin assignments identified using stable hydrogen isotopes in feathers (δ²H). We found support for spatial differentiation among routes by American redstarts and ovenbirds and temporal differentiation along routes by American redstarts. After controlling for breeding origin, the timing of American redstart migration differed among ages and sexes and ovenbird migration timing was influenced by carry‐over from winter habitat. The strength of migratory connectivity did not differ among the three species, with each showing weak breeding‐to‐spring migration MC relative to prior assessments of breeding‐wintering connectivity. Our work begins to fill an essential gap in methodology and understanding of the extent to which populations remain together during migration, information critical for a full annual cycle perspective on the population dynamics and conservation of migratory animals.     

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

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

Variation in climate warming along the migration route uncouples arrival and breeding dates

Migratory species are of special concern in the face of global climate change, since they may be affected by changes in the wintering area, along the migration route and at the breeding grounds. Here we show that migration and breeding times of a trans‐Saharan migrant, the pied flycatcher Ficedula hypoleuca, closely follow local temperatures along the migration route and at the breeding grounds. Because of differences in long‐term temperature trends of short within‐spring periods, the migration period and the time interval between migration and breeding dates of this species have extended in SW Finland. Temperatures in northern parts of Central Europe have risen at the time when the first migrants arrive there, facilitating their migration northward. Temperatures later in the spring have not changed, and the last individuals arrive at the same time as before. The timing of breeding has not advanced because temperatures at the breeding site after arrival have not changed. These results show that the pied flycatchers can speed up their migration in response to rising temperatures along the migration route. Our results strongly indicate that the effects of climate change have to be studied at the appropriate time and geographical scales for each species and population concerned.     

Body mass of six long-distance migrant passerine species along the autumn migration route

Summary We analysed body mass and moult data of six passerine species along their autumn migration route from northern Europe to North Africa and derived hypothetical models of the organisation of their migration in terms of fuel store accumulation. We analysed data of 46,541 first-year birds from 34 trapping sites, sampled in a network of collaborating European and African ringing stations. After accounting for effects of time of day and size, there were marked differences between the six species examined in the change of body mass along the migration route and in the timing of moult. Garden Warblers (Sylvia borin) and Pied Flycatchers (Ficedula hypoleuca) underwent their postjuvenile moult prior to migration and increased their average body mass along the migration route. Sedge Warblers (Acrocephalus schoenobaenus) also increased body mass towards the south, but started the migration bout without further refuelling well before the Sahara and moulted mainly in the wintering grounds. Reed Warblers (Acrocephalus scirpaceus) and Whitethroats (Sylvia communis) migrated while still moulting and did not increase average body mass towards south. They accumulated the energy needed to fly over the Sahara just before it. Spotted Flycatchers (Muscicapa striata) behaved in the same way, but contrary to Reed Warblers and Whitethroats they did not accumulate much fat stores in North Africa, which might urge them to stop and fuel up regularly in the Sahara. In the course of the season average body mass of all species increased slightly, which enabled them to migrate faster. In general, average body mass of first-year birds in northern and central Europe during the migration period was comparable to that of adults during breeding.

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Körpermassen von sechs Arten von Langstreckenziehern entlang des Herbstzugweges

Zusammenfassung Wir analysierten Körpermassen- und Mauserdaten von sechs Singvogelarten auf dem Herbstzug entlang ihres Zugweges von Nordeuropa bis Nordafrika. Die Körpermassen von 46 541 diesjährigen Vögeln, die auf 34 Beringungsstationen im Rahmen eines internationalen Projektes gesammelt wurden, sind die Grundlage dieser Untersuchung. Neben der Entwicklung der Körpermassen entlang des Zugweges untersuchten wir den Einfluss der Mauser, des Fangdatums, der Tageszeit und der Körpergröße auf die Körpermasse. Mausernde Vögel waren schwerer als nichtmausernde mit der gleichen Fettmenge. Die Körpermasse nahm im Laufe der Saison und im Verlauf des Tages zu. Große Vögel (mit einer längeren 8. Handschwinge) waren schwerer als kleine. Gartengrasmücken (Sylvia borin) und Trauerschnäpper (Ficedula hypoleuca) wurden entlang des Zugweges kontinuierlich schwerer. Beide Arten machten die Jugendteilmauser hauptsächlich noch im Brutgebiet durch. Schilfrohrsänger (Acrocephalus schoenobaenus) wurden nach Süden ebenfalls etwas schwerer, aber sie begannen mit einem Reiseabschnitt ohne weiteren Fettaufbau schon in Mitteleuropa und mauserten ihr Kleingefieder hauptsächlich erst in Afrika. Die Körpermassen von Teichrohrsängern (Acrocephalus scirpaceus) und Dorngrasmücken (Sylvia communis) blieben nach Süden weitgehend konstant. Beide Arten lagerten die nötigen Energiereserven zum Überfliegen der Sahara erst kurz davor an. Sie trennten Jugendteilmauser und Zug nicht vollständig. Ähnlich verhielten sich Grauschnäpper (Muscicapa striata), doch im Gegensatz zu den zwei vorherigen Arten, wurden sie auch im Mittelmeergebiet nicht wesentlich fetter. Sie scheinen daher regelmässig in der Wüste zu rasten und zu fressen.

     

Rapid formation of new migration route and breeding area by Arctic geese

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Extrapair paternity, migration, and breeding synchrony in birds

To understand interspecific patterns in the strength of sexualselection, variation in the costs and benefits of exercisingmate choice needs to be evaluated. One manifestation of sexualselection in birds is the occurrence of greatly variable levelsof extrapair paternity (EPP). A proposed general explanationfor this variation is that the benefits to females in seekingextrapair copulations vary in a predictable manner accordingto the degree of breeding synchrony because females are betterable to assess potential extrapair partners when males are simultaneouslyin breeding condition. This hypothesis predicts a latitudinaltrend in EPP because birds tend to breed more synchronouslyaway from the equator. Expanding on previous geographicallyand taxonomically restricted tropical/temperate comparisons,we used phylogenetically independent standardized linear contraststo show that this positive relationship persists when all birdspecies for which EPP estimates currently exist are considered.However, if a third factor covaries with latitude in the sameway as breeding synchrony and EPP, this relationship need notbe causal. Migration could also account for latitudinal variationin the benefits to females of pursuing EPP, if migration isassociated with (1) hasty or (2) inaccurate mate choice, (3)facilitated assessment of male quality through the condition-dependenceof arrival time, or (4) increased genetic variance in male quality.We show that migration distance is positively related to theproportion of EPP and that migration can statistically explainthe latitudinal trend in EPP, even when confounding factorsare simultaneously controlled. Hence, alternative explanationsfor latitudinal variation in EPP may be feasible, and carefulintraspecific tests are needed to assess their relative importanceand their implications for geographical variation in life-historyevolution.     

Is there a connection between weather at departure sites,onset of migration and timing of soaring‐bird autumn migration in Israel?

Aims Different aspects of soaring‐bird migration are influenced by weather. However, the relationship between weather and the onset of soaring‐bird migration, particularly in autumn, is not clear. Although long‐term migration counts are often unavailable near the breeding areas of many soaring birds in the western Palaearctic, soaring‐bird migration has been systematically monitored in Israel, a region where populations from large geographical areas converge. This study tests several fundamental hypotheses regarding the onset of migration and explores the connection between weather, migration onset and arrival at a distant site. Location Globally gridded meteorological data from the breeding areas in north‐eastern Europe were used as predictive variables in relation to the arrival of soaring migrants in Israel. Methods Inverse modelling was used to study the temporal and spatial influence of weather on initiation of migration based on autumn soaring‐bird migration counts in Israel. Numerous combinations of migration duration and temporal influence of meteorological variables (temperature, sea‐level pressure and precipitable water) were tested with different models for meteorological sensitivity. Results The day of arrival in Israel of white storks, honey buzzards, Levant sparrowhawks and lesser spotted eagles was significantly and strongly related to meteorological conditions in the breeding area days or even weeks before arrival in Israel. The cumulative number of days or cumulative value above or below a meteorological threshold performed significantly better than other models tested. Models provided reliable estimates of migration duration for each species. Main conclusions The meteorological triggers of migration at the breeding grounds differed between species and were related to deteriorating living conditions and deteriorating migratory flight conditions. Soaring birds are sensitive to meteorological triggers at the same period every year and their temporal response to weather appears to be constrained by their annual routine.     

Genetic variation reveals large-scale population expansion and migration during the expansion of Bantu-speaking peoples

The majority of sub-Saharan Africans today speak a number of closely related languages collectively referred to as ‘Bantu’ languages. The current distribution of Bantu-speaking populations has been found to largely be a consequence of the movement of people rather than a diffusion of language alone. Linguistic and single marker genetic studies have generated various hypotheses regarding the timing and the routes of the Bantu expansion, but these hypotheses have not been thoroughly investigated. In this study, we re-analysed microsatellite markers typed for large number of African populations that—owing to their fast mutation rates—capture signatures of recent population history. We confirm the spread of west African people across most of sub-Saharan Africa and estimated the expansion of Bantu-speaking groups, using a Bayesian approach, to around 5600 years ago. We tested four different divergence models for Bantu-speaking populations with a distribution comprising three geographical regions in Africa. We found that the most likely model for the movement of the eastern branch of Bantu-speakers involves migration of Bantu-speaking groups to the east followed by migration to the south. This model, however, is only marginally more likely than other models, which might indicate direct movement from the west and/or significant gene flow with the western Branch of Bantu-speakers. Our study use multi-loci genetic data to explicitly investigate the timing and mode of the Bantu expansion and it demonstrates that west African groups rapidly expanded both in numbers and over a large geographical area, affirming the fact that the Bantu expansion was one of the most dramatic demographic events in human history.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

Extremely detoured migration in an inexperienced bird: interplay of transport costs and social interactions

We tagged two juvenile short‐toed eagles in southern Italian peninsula with GPS satellite transmitters. According to previous visual observations, two different migratory routes for Italian short‐toed eagles to reach Africa in autumn have been proposed: via Sicily and via Gibraltar. These routes include different over‐water distances to cross the Mediterranean Sea, and thus different proportions of flight modes (soaring–gliding vs flapping–gliding) with resulting different transport costs. Considering different scenarios of energy cost of transport, with flapping–gliding flight over water being more costly than flying over land using soaring–gliding flight, we predicted a maximum optimal detour of 1218 km. Both individuals reached Africa using the longest, detoured, route, avoiding the longest water crossing. To achieve this they began migrating northwards, keeping for ca 700 km a direction opposite to that followed by any other migrating bird from the Northern hemisphere in autumn. The comparison of optimal detour predictions with observed migratory tracks suggests that this migratory strategy prioritizes not only energy minimization, but also safety, given the mortality risk associated with the sea crossing. Finally, it is unlike that these inexperienced individuals followed such a complex route relying only on endogenous information and we therefore suggest, also on the basis of field observations, that social interactions (adult guidance) allow these individuals to learn the detoured route.     

Avian migrants adjust migration in response to environmental conditions en route

The onset of migration in birds is assumed to be primarily under endogenous control in long-distance migrants. Recently, climate changes appear to have been driving a rapid change in breeding area arrival. However, little is known about the climatic factors affecting migratory birds during the migration cycle, or whether recently reported phenological changes are caused by plastic behavioural responses or evolutionary change. Here, we investigate how environmental conditions in the wintering areas as well as en route towards breeding areas affect timing of migration. Using data from 1984 to 2004 covering the entire migration period every year from observatories located in the Middle East and northern Europe, we show that passage of the Sahara Desert is delayed and correlated with improved conditions in the wintering areas. By contrast, migrants travel more rapidly through Europe, and adjust their breeding area arrival time in response to improved environmental conditions en route. Previous studies have reported opposing results from a different migration route through the Mediterranean region (Italy). We argue that the simplest explanation for different phenological patterns at different latitudes and between migratory routes appears to be phenotypic responses to spatial variability in conditions en route.     

Fuel deposition of three passerine bird species along the migration route

The rate at which migrant birds replenish their energy stores at intermittent stopovers largely determines overall migration speed, the manner in which migration proceeds and success of migration. In this study, data on the fuel deposition rate (FDR) of three long-distance migrants from 17 ringing sites along their autumn migration route were used to examine: (1) effects of endogenous factors on FDR, and (2) how relationships between exogenous factors and FDR affect the organisation of migration. We developed a model to estimate FDR from retrapped birds which takes into account time of day and various other factors which might influence FDR. The two endogenous factors, moult and current energy stores, generally reduced FDR. This may result in lower departure energy loads and more stopovers than expected from optimal migration theory. Differences between species with respect to seasonal, year-to-year and geographical patterns of FDR could be related to differences in availability and predictability of food resources, and help to explain differences in the organisation of migration. A low FDR in northern and central Europe could be related to low, but predictable, food resources and an early departure during moult of the reed warbler (Acrocephalus scirpaceus); FDRs varying between years were related to large spatial and year-to-year variation in the density of the main prey of the sedge warbler (Acrocephalus schoenobaenus); and a high FDR in the garden warbler (Sylvia borin) was related to abundant food resources, due to a switch from a purely invertebrate diet to a mixed diet including fruits which are abundant over large areas of Europe and north Africa. This study demonstrated that the organisation of migration is the outcome of a complex interplay of the seasonal timing of moult, food availability and predictability and a seasonal switch in diet, and can be modified by individual birds in response to a limited amount of time in which to migrate. Received: 26 April 1999 / Accepted: 24 September 1999     

Partial migration in inexperienced pied avocets Recurvirostra avosetta: distribution pattern and correlates

Birds exhibit a range of wintering behaviour from strictly migrant to strictly resident species. In partially migrating ones, some birds overwinter within their breeding region (resident birds) while others, although breeding in the same area, winter far away (migrant birds). Accordingly, choosing a wintering region is a key stage in the annual life cycle of birds, notably for inexperienced first‐year individuals. The present study aimed to investigate this issue, and more specifically to study the distribution pattern during winter and factors influencing the wintering behaviour of first‐year pied avocets Recurvirostra avosetta. Based on a 10‐yr ringing study carried out on five of the major French breeding colonies distributed along the Atlantic coast, we showed the coexistence of different wintering tactics. The resident tactic was predominant (approximately 86% of the 575 birds re‐sighted), while the other birds adopted migration. Among resident individuals, two different tactics occurred: 43% of birds overwintered within their natal colony, whereas the others wintered in another site located at relatively close proximity along the French Atlantic coast. Hatching date was a consistent predictor of all wintering tactics. More specifically, the probability of migrating was the highest for early‐hatched birds, and for resident ones, the probability of wintering within their natal colony rather than in another French site was the highest for both median‐ and late‐hatched individuals. In addition, a colony effect was demonstrated for resident birds. Several biological interpretations, including social system, variations in both individual body condition and habitat quality, were put forward to explain these correlates.     

Effect of carbon dioxide on the Oriental cockroach Blatta orientalis

Exposure to high concentrations of carbon dioxide (CO2) killed adult and nymphal stages of the oriental cockroach Blatta orientalis L. (Dictyoptera: Blattidae) with LT50 values of 11.5-16.2 h for 60% CO2 in air and 5.7-7.1 h for 100% CO2 at 20 degrees C; corresponding LT50s at 28 degrees C were 2.8-4.6 h for 60% CO2 in air and 2.3-3.6 h for 100% CO2. Complete kill of mobile stages was obtained within 24 h using 60% CO2 at 20 degrees C. Survivors of treatments with 100% CO2 at 28 degrees C remained completely paralysed for up to 3 days post-treatment and took up to 5 days to regain normal movement, but adult females then resumed production of oothecae with no significant loss in fecundity. Oothecae 5 or 30 days after deposition required 60-84 h exposure to 60% CO2 at 20 degrees C to prevent emergence of nymphs but less time using 100% CO2 at 28 degrees C. At 28 degrees C, when adult females were treated with 100% CO2 and 52% r.h. for 6 h (giving 100% mortality) loss of weight was significantly greater than that following treatment with air at 52% r.h. for 6 h (giving no mortality). However, significantly greater weight loss also occurred when they were treated with dried air (< 10% r.h.) for 6 h, also with no mortality. The toxicity of CO2 to mobile stages of the oriental cockroach appeared to result from irreversible effects on the nervous system, rather than from water loss during exposure.     

Non‐breeding areas and timing of migration in relation to weather of Scottish‐breeding common sandpipers Actitis hypoleucos

The number of breeding common sandpipers has declined in Britain due to poorer return rates from non‐breeding areas. To investigate little known aspects of their annual cycle, breeding common sandpipers were fitted with geolocators to track their migrations and determine their non‐breeding areas. Ten tagged birds left Scotland on 9 July (median dates and durations are given throughout the abstract). Short‐term staging was carried out by some birds in England and Ireland, then for longer by most birds in Iberia before continuing to West Africa, arriving on 28 July. Six birds spent most of the non‐breeding season (October–February) on the coast of Guinea‐Bissau, suggesting that this is a key area. Single birds occurred in Sierra Leone, Guinea, the Canary Islands and western Sahara. The southward migration from Scotland took 17.5 d (range 1.5–24 d), excluding the initial fuelling period. The first northward movement from Africa was on 12 April. Staging occurred in either Morocco, Iberia or France. Arrival in Scotland was on 2 May. The northward migration took 16 d (range 13.5–20.5 d). The main migration strategy involved short‐ and medium‐range flights, using tail‐winds in most cases. Variation in strategy was associated with departure date; birds that left later staged for shorter durations. Coastal West Africa provides two major habitats for common sandpipers: mudflats associated with mangroves and rice fields. Although the area of mangrove has been depleted, the scale of loss has probably been insufficient to account for the decline in sandpiper numbers. Rice fields are expanding, providing feeding areas for water‐birds. Meteorological data during the migrations suggest that the weather during the southward migration is unlikely to contribute to a population decline but strong cross‐winds or head‐winds during the northward migration to the breeding grounds may do so.     

Biological and Chemical Induction of Resistance to the Globodera tabacum solanacearum in Oriental and Flue-Cured Tobacco (Nicotiana tabacum L.)

The effects of acibenzolar-S-methyl (ASM) and four combinations of plant growth-promoting rhizobacteria (PGPR) on the reproduction of a tobacco cyst nematode, Globodera tabacum solanacearum, and growth of Nicotiana tabacum (cv. K326 and Xanthi) were tested under greenhouse and field conditions. The PGPR included combinations of Bacillus subtilis A13 with B. pumilis INR7, B. pumilis SE34, B. licheniformis IN937b, or B. amyloliquefaciens IN937a, respectively. Among the four rhizobacterial combinations, IN937a + A13 exhibited the most consistent reduction in G. t. solanacearum cysts under greenhouse and field conditions. No undesirable effects of IN937a + A13 were observed on tobacco growth under greenhouse and field conditions. Use of INR7 + A13 reduced G. t. solanacearum reproduction on flue-cured tobacco cv. K326 but not on oriental tobacco cv. Xanthi. Application of ASM reduced final numbers of G. t. solanacearum cysts, but also resulted in phytotoxicity mainly under the greenhouse conditions. When oriental tobacco seedlings were pre-grown in a IN937a + A13-treated soil-less medium, a single application of ASM at 200 mg/L one week after transplanting significantly reduced G. t. solanacearum reproduction in the field.