Zugunruhe of migratory and non-migratory birds in a circannual context

Recent findings in non-migratory birds have reopened questions about the interpretation and seasonal organization of Zugunruhe . I address the relationship between Zugunruhe and migration by comparing underlying circannual patterns in captive populations of migratory and non-migratory stonechats. Zugunruhe was highly variable and lacked clear periodicity, indicating its sensitivity to external cues. Patterns of Zugunruhe were similar in African residents, European short-distance migrants, and Siberian long-distance migrants, revealing no major difference in circannual organization. Moult was regulated independently of Zugunruhe and timed more rigidly, particularly in stonechats from equatorial Africa. The persistent and variable circannual patterns of Zugunruhe suggest that non-migratory and migratory stonechats have similar underlying programs but have modified the expression of actual migration. The findings, together with published observations from other species, emphasize the importance of considering programs for migration in a wide range of species, without losing sight of its environmental context.     

Nocturnal Activity of Caged Resident and Migrant Silvereyes (Zosteropidae: Aves)

Night-activity patterns of individually-caged grey-breasted silvereyes Zosterops lateralis of resident and migrant status were investigated. True migratory restlessness or ‘Zugunruhe’ was exhibited by some individuals, but there was also considerable ‘noise’ in the data caused by cageing and neighbouring disturbance. In general, ‘Zugunruhe’ took place from about 2 h before, to about 4 h after ‘sunrise’, and both the daily timing and annual duration of Zugunruhe were consistent with the migration pattern observed in the field. A total of 42 % of resident birds from Tasmania, 77 % of migrant birds from Tasmania and 90 % of migrant birds from the mainland were considered to have exhibited true ‘Zugunruhe’, while none of the resident birds from Brisbane were considered to have displayed it. A few individuals spontaneously developed ‘Zugunruhe’ in the absence of other birds, but many birds (both residents and migrants) displayed no elevated night unrest when placed in a room on their own, although displaying unrest when placed among birds exhibiting ‘Zugunruhe’. The migration of the silvereye probably has a genetic basis, but whether an individual migrates or not is also influenced by social factors which differ in magnitude between individuals.     

Size‐dependent costs of migration: Migrant bird species are subordinate to residents,but only at small body sizes

Migrant species are commonly thought to be poor competitors in aggressive interactions with resident species. However, no studies have tested whether this relationship is widespread. Here, we compare the behavioural dominance of closely related species of migratory and nonmigratory birds, testing whether migrants are consistently subordinate to resident species in aggressive contests. We compiled published behavioural dominance data involving migrant and resident congeners, gathering additional data on the body mass and migratory distance of each species. Focal species included a diverse array of birds (28 taxonomic families, 12 orders) from around the world. We found that migrant species are usually subordinate to resident species, but that this relationship disappears at larger body sizes. For smaller birds (<500 g), resident species were behaviourally dominant in 83%–88% of comparisons; for larger birds (>500 g), resident species were dominant in only 25%–30% of comparisons. The relative difference in body mass best predicted dominance relationships among species, with larger species dominant in 80%–84% of comparisons. When migrant and resident masses were equal, however, resident species were still more likely to be dominant in smaller birds, suggesting that other factors may also contribute to the subordinate status of migrants. Overall, our results suggest that in smaller species, the evolution of migration is associated with lighter weights and other traits that compromise the competitive abilities of migrants relative to residents. In contrast, larger species appear able to evolve migration without compromising their size or competitive abilities in aggressive contests, suggesting size‐dependent constraints on the evolution of migration.     

Migratory sleeplessness in the white-crowned sparrow (Zonotrichia leucophrys gambelii)

Twice a year, normally diurnal songbirds engage in long-distance nocturnal migrations between their wintering and breeding grounds. If and how songbirds sleep during these periods of increased activity has remained a mystery. We used a combination of electrophysiological recording and neurobehavioral testing to characterize seasonal changes in sleep and cognition in captive white-crowned sparrows (Zonotrichia leucophrys gambelii) across nonmigratory and migratory seasons. Compared to sparrows in a nonmigratory state, migratory sparrows spent approximately two-thirds less time sleeping. Despite reducing sleep during migration, accuracy and responding on a repeated-acquisition task remained at a high level in sparrows in a migratory state. This resistance to sleep loss during the prolonged migratory season is in direct contrast to the decline in accuracy and responding observed following as little as one night of experimenter-induced sleep restriction in the same birds during the nonmigratory season. Our results suggest that despite being adversely affected by sleep loss during the nonmigratory season, songbirds exhibit an unprecedented capacity to reduce sleep during migration for long periods of time without associated deficits in cognitive function. Understanding the mechanisms that mediate migratory sleeplessness may provide insights into the etiology of changes in sleep and behavior in seasonal mood disorders, as well as into the functions of sleep itself.     

Migratory Sleeplessness in the White-Crowned Sparrow (Zonotrichia leucophrys gambelii)

Twice a year, normally diurnal songbirds engage in long-distance nocturnal migrations between their wintering and breeding grounds. If and how songbirds sleep during these periods of increased activity has remained a mystery. We used a combination of electrophysiological recording and neurobehavioral testing to characterize seasonal changes in sleep and cognition in captive white-crowned sparrows (Zonotrichia leucophrys gambelii) across nonmigratory and migratory seasons. Compared to sparrows in a nonmigratory state, migratory sparrows spent approximately two-thirds less time sleeping. Despite reducing sleep during migration, accuracy and responding on a repeated-acquisition task remained at a high level in sparrows in a migratory state. This resistance to sleep loss during the prolonged migratory season is in direct contrast to the decline in accuracy and responding observed following as little as one night of experimenter-induced sleep restriction in the same birds during the nonmigratory season. Our results suggest that despite being adversely affected by sleep loss during the nonmigratory season, songbirds exhibit an unprecedented capacity to reduce sleep during migration for long periods of time without associated deficits in cognitive function. Understanding the mechanisms that mediate migratory sleeplessness may provide insights into the etiology of changes in sleep and behavior in seasonal mood disorders, as well as into the functions of sleep itself.     

Diet of breeding Eleonora's falcon Falco eleonorae in Algeria: Insights for the autumn trans‐Mediterranean avian migration

How environmental changes are affecting bird population dynamics is one of the most challenging conservation issues. Dietary studies of top avian predators could offer scope to monitor anthropogenic drivers of ecosystem changes. We investigated the diet of breeding Eleonora''s falcon in an area of Northeastern Algeria in the years 2010–2012. Feathers and insect remains originating from prey plucking behavior were analyzed, providing insights into the seasonally changing diet of this raptor, as well as the trans‐Mediterranean avian migration. A total of 77 species of birds (16 Sylviidae, 11 Turdidae, and 4 Emberizidae), 3 species of insects, and 1 lizard were identified among prey remains, reflecting a diverse diet. Diet composition and prey abundance varied seasonally, faithfully correlating with the passage of migrant birds as recorded from bird ring recoveries. Our findings suggest that dietary studies of predators might be deployed to investigate changes in bird migration. We discuss our results in the context of trans‐Mediterranean migration, with early‐season prey mainly comprising trans‐Saharan migrants (Apus apus and Merops apiaster) and late‐season prey being dominated by Mediterranean winter migrants (Erithacus rubecula, Turdus philomelos, Sylvia atricapilla, and Sturnus vulgaris). Notably, we observed a significant reduction in species richness of passerine remains in 2012, potentially highlighting a decline in the diversity of avian migrants.     

The actin regulators Enabled and Diaphanous direct distinct protrusive behaviors in different tissues during Drosophila development

Actin-based protrusions are important for signaling and migration during development and homeostasis. Defining how different tissues in vivo craft diverse protrusive behaviors using the same genomic toolkit of actin regulators is a current challenge. The actin elongation factors Diaphanous and Enabled both promote barbed-end actin polymerization and can stimulate filopodia in cultured cells. However, redundancy in mammals and Diaphanous’ role in cytokinesis limited analysis of whether and how they regulate protrusions during development. We used two tissues driving Drosophila dorsal closure—migratory leading-edge (LE) and nonmigratory amnioserosal (AS) cells—as models to define how cells shape distinct protrusions during morphogenesis. We found that nonmigratory AS cells produce filopodia that are morphologically and dynamically distinct from those of LE cells. We hypothesized that differing Enabled and/or Diaphanous activity drives these differences. Combining gain- and loss-of-function with quantitative approaches revealed that Diaphanous and Enabled each regulate filopodial behavior in vivo and defined a quantitative “fingerprint”—the protrusive profile—which our data suggest is characteristic of each actin regulator. Our data suggest that LE protrusiveness is primarily Enabled driven, whereas Diaphanous plays the primary role in the AS, and reveal each has roles in dorsal closure, but its robustness ensures timely completion in their absence.     

Vergleichende Untersuchung von Jugendentwicklung und Zugverhalten bei Garten- und Hausrotschwanz,Phoenicurus phoenicurus undPh. ochruros

Zusammenfassung An je 12 handaufgezogenen Garten- und Hausrotschwänzen wurden Jugendmauser, Fettdeposition und Zugunruhe erfaßt und mit entsprechenden Daten freilebender Artgenossen verglichen. Die Daten stimmen gut mit denen freilebender Artgenossen überein. Beim Gartenrotschwanz setzt die Jugendmauser in frühem Alter ein und verläuft rasch, und Fettdeposition und Zugunruhe beginnen, ehe die Mauser beendet wird. Er zeigt das typische Bild eines frühwegziehenden Interkontinentalziehers. Beim Hausrotschwanz hingegen wird die Jugendmauser trotz ihrer großen Länge beendet, bevor Fettdeposition und Zugunruhe einsetzen, und beide haben geringeren Umfang. Er bietet das Bild eines weniger ausgeprägten Ziehers und eines Intrakontinentalziehers. Beide Arten zeigen ihre großen Unterschiede in Jugendentwicklung und Zugverhalten in weitgehend entsprechenden Versuchsbedingungen, vor allem unter ganz ähnlichen photoperiodischen Bedingungen. Das spricht dafür, daß diese Unterschiede einer stark endogenen, wahrscheinlich einer stark genetischen Kontrolle unterliegen. Der Gartenrotschwanz — als Weitstrekkenzieher — entwickelt viel, der Hausrotschwanz — als Kurz- bis Mittelstreckenzieher — weit weniger Nachtunruhe. Die Zeit, in der Nachtunruhe produziert wird, stimmt bei beiden Arten sehr gut mit der Zugzeit überein. Die Nachtunruhe ist daher bei beiden Arten typische Zugunruhe. Die verschiedenen gemessenen Zugunruhewerte stimmen z. T. sehr gut mit den zurückzulegenden Zugstrecken überein, z. T. zeigen sie erhebliche Abweichungen. Prinzipiell lassen sich die Daten jedoch alle gut im Sinne endogener Zug-Zeitprogramme interpretieren, wie sie früher für Sylviiden nachgewiesen wurden. Demnach lassen sich die Vorstellungen endogener Zug-Zeitprogramme nunmehr auf eine weitere Vogelgruppe, nämlich die kleinen Drosselvögel, ausdehnen.

---

A comparative study of juvenile development, migratory restlessness and migratory behaviour in the Redstart and Black Redstart

Summary Juvenile development, migratory disposition and migratory activity were investigated in Redstarts and Black Redstarts. In each 12 hand-raised individuals, patterns of juvenile moult, fat deposition and migratory restlessness were determined and compared with corresponding data from freeliving conspecifics. There was good agreement between the data from hand-raised and feral birds. In the Redstart, juvenile moult starts early and proceeds rapidly and fat deposition and migratory restlessness start before the moult has ended. The Redstart is typical for an early departing intercontinental migrant. In the Black Redstart, however, the moult, despite its long duration was clearly separated from the migratory events, and fat deposition and migratory restlessness are both less pronounced than in the Redstart. The patterns of the Black Redstart were typical for a less pronounced or an intracontinental migrant. Both species produced the large differences in juvenile development and migratory behaviour in widely corresponding experimental conditions, above all in very similar photoperiodic conditions. This suggests that these differences were based to a high degree on endogenous, probably genetic factors. The Redstart, as a long-distance migrant, developed a lot, and the Black Redstart, a short- to middle-distance migrant, far less nocturnal restlessness. In both species the period in which nocturnal restlessness developed corresponded very well with the migratory season. Thus this restlessness was typical migratory restlessness. The various measurements of restlessness obtained corresponded in part with the distances covered during migration although there were substantial deviations. In principle, however, all the data can well be interpreted in terms of endogenous time programs for migration as they have been formerly demonstrated for Sylviid species. Thus the concept of these programs can now be extended to another bird group — to small Turdiid species.

     

A separate circadian oscillator controls nocturnal migratory restlessness in the songbird Sylvia borin

When confined to a cage, migratory songbirds exhibit nocturnal migratory restlessness (also called Zugunruhe) during the spring and autumn migratory periods, even though these birds are exclusively diurnal during the remainder of the year. Zugunruhe, which has been demonstrated to be under the direct control of a circannual timer, is characterized by a stereotypic "wing-whirring" behavior while the bird is perched. To elucidate the role played by the circadian system in the regulation of Zugunruhe, the authors studied the activity of garden warblers (Sylvia borin), long-distance nocturnal migrants, under skeleton photoperiods of different lengths and under constant dim light. In 11.5D:1L:10.5D:1L skeleton photoperiods, the authors found that Zugunruhe free-ran in a substantial proportion of birds, while their normal daily activities (e.g., feeding and preening) remained synchronized to 24 h. Some birds expressing Zugunruhe under constant dim light continued to show 2 distinct bouts of activity: one corresponding to daily activities, the other to wing-whirring. In some cases, these 2 bouts crossed while free-running with different periods. Birds expressing Zugunruhe also had significantly longer free-running periods than birds that did not. The study data suggest that the seasonal appearance of Zugunruhe is the result of the interactions of at least 2 circadian oscillators and that it is the phase relationship of these 2 oscillators that determines when nocturnal migratory restlessness is expressed. Furthermore, these data are consistent with the previously proposed internal coincidence hypothesis as a model for the ontogeny of circannual rhythms.     

Size and accumulation of fuel reserves at stopover predict nocturnal restlessness in a migratory bird

Early arrival at the breeding site positively affects the breeding success of migratory birds. During migration, birds spend most of their time at stopovers. Therefore, determining which factors shape stopover duration is essential to our understanding of avian migration. Because the main purpose of stopover is to accumulate fat as fuel for the next flight bout, fuel reserves at arrival and the accumulation of fuel are both expected to affect stopover departure decisions. Here, we determined whether fuel reserves and fuel accumulation predict a bird''s motivation to depart, as quantified by nocturnal migratory restlessness (Zugunruhe), using northern wheatears (Oenanthe oenanthe) that were captured and temporarily contained at spring stopover. We found that fuel reserves at capture were positively correlated with Zugunruhe, and negatively correlated with fuel accumulation. This indicates that fat birds were motivated to depart, whereas lean birds were set on staying and accumulating fuel. Moreover, the change in fuel reserves was positively correlated with the concurrent change in Zugunruhe, providing the first empirical evidence for a direct link between fuel accumulation and Zugunruhe during stopover. Our study indicates that, together with innate rhythms and weather, the size and accumulation of fuel reserves shape stopover duration, and hence overall migration time.     

Food availability but not melatonin affects nocturnal restlessness in a wild migrating passerine

A large number of passerine species migrate at night, although most of them are diurnal outside the migratory seasons. This diurnal-to-nocturnal transition is a major life-history event, yet little is known about its physiological control. Previous work showed that during the migratory periods captive birds showing nocturnal migratory restlessness (Zugunruhe) have reduced concentrations of circulating melatonin at night compared to non-migratory periods. This suggested that the hormone melatonin, a main component of the avian circadian system, is involved in the expression of Zugunruhe. Other studies demonstrated that the relationship between low melatonin levels and Zugunruhe is not a seasonal correlation. When Zugunruhe was interrupted by exposing birds to a fasting-and-refeeding protocol, melatonin levels increased. Here we studied whether melatonin and food availability influence the intensity of Zugunruhe in wild migrating garden warblers (Sylvia borin) at a stopover site. Birds were held in recording cages overnight, with or without food available, and either bled to determine melatonin concentrations or treated transdermally with melatonin. We found that melatonin levels at night were correlated with the intensity of diurnal locomotor activity and with condition, but were not correlated with Zugunruhe. Similarly, the melatonin treatment did not have effects on Zugunruhe, whereas food availability increased it. Our study shows that the nocturnal melatonin levels in migrating warblers depend on food availability and are correlated with condition. In addition, it suggests that melatonin does not control Zugunruhe and might rather be involved in energy conservation and/or clock synchronization during migration.     

Carry-over effects of day length during spring migration

The day lengths to which migratory birds are exposed depend on the timing and course of their journey. While winter day length is known to influence vernal events, it is not clear if birds also use day length during the spring migration as a temporal cue. We addressed this question by exposing captive stonechats (Saxicola torquata) to two different photoperiodic simulations of spring migration routes, following common winter conditions. One group experienced day lengths of the regular (“fast”) migration, and the other group, a “slow”, or more southerly originating, route. The resulting small, temporary differences in day length had lasting effects on the birds. The groups differed in migratory restlessness during and following exposure to different day lengths. “Slow” migrants continued nocturnal activity longer than “fast” migrants. Furthermore, all activities of the ensuing breeding season were delayed in the “slow” migrants, indicating a phase shift in their underlying annual rhythm. “Slow” migrants delayed terminating their reproductive stage by regressing testes and the cloacal protuberance later than the “fast” migrants. Molt started and ended later in “slow” migrants, but the duration of the molt was unaffected by spring day length. Finally, “fast” migrants resumed nightly restlessness earlier than “slow” migrants in late summer. These results demonstrate that Zugunruhe (migratory restlessness) and reproductive windows are not set exclusively during winter but can be modified by day length cues during the spring migration. Because migration modifies the day length exposure of birds, migration routes can have carry-over effects on the timing of breeding season events, including the completion of molt and initiation of autumnal nocturnal activity.

---

Einfluss der Tagesl?nge w?hrend des Frühjahrszugs auf Brutzeitraum, Mauser, und Beginn der Herbstzugunruhe

Zusammenfassung Zugv?gel beeinflussen durch ihre Zugroute und Zugzeit die Tagesl?nge, in der sie sich befinden. W?hrend experimentell belegt ist, dass die Wintertagesl?nge das saisonale Verhalten im Frühling beeinflusst, ist bisher unbekannt, ob V?gel auch w?hrend der Frühjahrszugzeit photoperiodische Zeitinformationen nutzen. Um diese Frage zu beantworten, haben wir Schwarzkehlchen (Saxicola torquata) unter den Lichtbedingungen von zwei verschiedenen Zugrouten untersucht. Die V?gel hatten unter einheitlichen Bedingungen überwintert. Anschlie?end erlebte eine Gruppe die Lichtbedingungen ihres regul?ren (“schnellen“) Zugweges, die andere die eines “langsameren“, oder weiter südlich beginnenden Zugweges. Die damit verbundenen geringen Unterschiede in der Tagesl?nge zeigten nachhaltige Wirkung. V?gel der beiden Gruppen unterschieden sich in Zugunruhe w?hrend und nachdem sie unterschiedliche Tagesl?ngen erlebten. “Langsame” Zieher setzten ihre n?chtliche Aktivit?t l?nger fort als “schnelle" Zieher. Darüber hinaus verz?gerten sich bei den “langsamen" Ziehern alle Aktivit?ten der anschlie?enden Brutsaison. Dies deutet auf eine Phasenverschiebung der zugrunde liegenden Jahresrhythmen hin. “Langsame" Zieher verz?gerten den Abschluss ihrer Brutphase, indem sie Hoden und Kloakenprotuberanz sp?ter zurückentwickelten als “schnelle" Zieher. Die Mauser begann und endete sp?ter als bei “schnellen" Ziehern, dauerte aber gleich lang an. Schlie?lich nahmen “schnelle" Zieher die n?chtliche Unruhe früher wieder auf als “langsame" Zieher. Diese Ergebnisse zeigen, dass der Brutzeitraum von Zugv?geln nicht allein im Winter, sondern auch w?hrend des Frühjahrszugs von der Tagesl?nge beeinflusst wird. Somit k?nnen sich Zugzeit und Zugroute auf das anschlie?ende zeitliche Verhalten bis zu Mauserende und Beginn der Herbstzugunruhe auswirken.

     

Temperature alters the photoperiodically controlled phenologies linked with migration and reproduction in a night-migratory songbird

We investigated the effects of temperature on photoperiodic induction of the phenologies linked with migration (body fattening and premigratory night-time restlessness, Zugunruhe) and reproduction (testicular maturation) in the migratory blackheaded bunting. Birds were exposed for four weeks to near-threshold photoperiods required to induce testicular growth (11.5 L:12.5 D and 12 L:12 D) or for 18 weeks to a long photoperiod (13 L:11 D) at 22°C or 27°C (low) and 35°C or 40°C (high) temperatures. A significant body fattening and half-maximal testicular growth occurred in birds under the 12 L, but not under the 11.5 L photoperiod. Further, one of six birds in both temperature groups on 11.5 L, and four and two of six birds, respectively, in low- and high-temperature groups on 12 L showed the Zugunruhe. Buntings on 13 L in both temperature groups showed complete growth-regression cycles in body fattening, Zugunruhe and testis maturation. In birds on 13 L, high temperature attenuated activity levels, delayed onset of Zugunruhe by about 12 days, reduced body fattening and slowed testicular maturation. The effect of temperature seems to be on the rate of photoperiodic induction rather than on the critical day length. It is suggested that a change in temperature could alter the timing of the development of phenologies linked with seasonal migration and reproduction in migratory songbirds.     

Simulation of migratory flight and stopover affects night levels of melatonin in a nocturnal migrant

Several species of diurnal birds are nocturnal migrants. The activation of nocturnal activity requires major physiological changes, which are essentially unknown. Previous work has shown that during migratory periods nocturnal migrants have reduced night-time levels of melatonin. Since this hormone is involved in the modulation of day-night rhythms, it is a good candidate regulator of nocturnal migratory activity. We studied whether melatonin levels change when nocturnally active blackcaps (Sylvia atricapilla) are experimentally transferred from a migratory to a non-migratory state. We simulated a long migratory flight by depriving birds of food for 2 days, and a refuelling stopover by subsequently re-administering food. Such a regimen is known to induce a reduction in migratory restlessness ('Zugunruhe') in the night following food reintroduction. The experiments were performed in both autumn and spring using blackcaps taken from their breeding grounds (Sweden) and their wintering areas (Kenya). In autumn, the food regimen induced a suppression of Zugunruhe and an increase in melatonin in the night following food reintroduction. In spring, the effects of the treatment were qualitatively similar but their extent depended on the amount of body-fat reserves. This work shows that the reduction of night-time melatonin during migratory periods is functionally related to nocturnal migration, and that fat reserves influence the response of the migratory programme to food deprivation.     

Migratory dark-eyed juncos, Junco hyemalis, have better spatial memory and denser hippocampal neurons than nonmigratory conspecifics

The evolution of migration in an animal population produces a suite of physiological, behavioural and cognitive adaptations. Migratory birds, in particular, require the ability to return annually to breeding and wintering sites after long journeys, and thus might be predicted to have evolved enhanced spatial memory. In a comparison of two sparrow subspecies that co-occur in winter, the migratory subspecies (dark-eyed junco, Junco hyemalis hyemalis) performed better than the nonmigratory subspecies (J. h. carolinensis) on a room-scale spatial memory test. The migratory juncos also had more densely packed hippocampal neurons than did nonmigrants. Among nonmigrants, we looked for hippocampal differences between birds that occupied two home ranges annually and those that remained on their breeding territory year-round, to determine whether migration, per se, is related to neuroanatomical differences. However, we were unable to reach any conclusions because of low statistical power. A denser hippocampus could be the basis for better spatial memory in migrant juncos. Further testing of spatial memory on a landscape scale is needed to strengthen this argument and to understand cognitive differences between migrants and nonmigrants. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

鸟类迁徙:在全球变暖趋势下的演化、调控与发展(英文)

最近几十年的研究证实 ,鸟类迁徙在很大程度上受到遗传因素的直接控制。有证据表明 ,存在某种先天的迁徙动因并涉及以下几方面的遗传调控 :(1)迁徙过程的起始、持续以及结束 ;(2 )迁徙活动量 ,即决定鸟类飞行距离的遗传参数 ;(3)迁徙方向 ;(4)生理参数 ,特别是迁徙期间的脂肪贮存 ,以及对于那些部分个体迁徙的鸟种而言 ,决定个体迁徙与否的生理参数。双因素选择实验表明 ,部分迁徙群经由几个世代的选择即可转变成完全的迁徙群或非迁徙群。新迁徙方向以及由此导致的新越冬区的改变 ,也能在野生鸟类中迅速实现。至少在以往研究得最为透彻的鸟种 (黑顶林莺Sylviaatricapilla)中 ,“迁徙”或“非迁徙”是先天性的 ,与特异性迁徙活动量相关 (尤如一时间程序 ) ,前者 (迁徙的 )已证实是由一种阈机制所控制的。一项新的鸟类迁徙理论假设 ,即使好些完全迁徙的类群 ,较低水平的迁徙活动量选择也会导致阈的异位 ,低于这一阈值就会出现非迁徙个体。因此 ,通过选择作用 ,一个迁徙型种群可以通过部分迁徙型转变为非迁徙型。这种中间阶段在现存鸟类中十分普遍。它始见于生物演化早期 ,就鸟类而言 ,可能在原始鸟类就已具备。模型运算表明 ,在施以强定向选择情况下 ,迁徙鸟类经过约 4 0年可转变为留鸟 ,反之亦然。这就解     

Stopover decision during migration: physiological conditions predict nocturnal restlessness in wild passerines

During migration, a number of bird species rely on stopover sites for resting and feeding before and after crossing ecological barriers such as deserts or seas. The duration of a stopover depends on the combined effects of environmental factors, endogenous programmes and physiological conditions. Previous studies indicated that lean birds prolong their refuelling stopover compared with fat birds; however, the quantitative relationship between physiological conditions and stopover behaviour has not been studied yet. Here, we tested in a large sample of free-living birds of three European passerines (whinchats, Saxicola rubetra, garden warblers, Sylvia borin and whitethroats, Sylvia communis) whether the amount of migratory restlessness (Zugunruhe) shown at a stopover site depends on physiological conditions. An integrated measure of condition based on body mass, amount of subcutaneous fat and thickness of pectoral muscles strongly predicted the intensity of Zugunruhe shown in recording cages in the night following capture. These results provide novel and robust quantitative evidence in support of the hypothesis that the amount of energy reserves plays a major role in determining the stopover duration in migratory birds.     

Spring passerine migrants stopping over in the Sahara are not fall-outs

The strategy of migrants crossing the Sahara desert has been the subject of debate, but recent evidence from radar studies has confirmed that most passerines use an intermittent migration strategy. The latter has also been suggested from previous studies in oases during autumn migration. It was found that migrants with relatively high fuel loads rest in the desert during daytime and continue migration during the following night, whereas lean migrants stopover in oases for several days to refuel. However, data from the Sahara are scarce for spring migration. We captured passerine migrants near B?r Amrane (22°47′N, 8°43′W) in the plain desert of Mauritania for 3 weeks during spring migration in 2004. We estimated flight ranges of 85 passerines stopping over in the desert to test whether they carried sufficient fuel loads to accomplish migration across the Sahara successfully. High fat loads of the majority of birds indicated that they were neither “fall-outs” nor too weak to accomplish migration successfully. The flight range estimates, based on mean flight speeds derived from radar measurements (59 km/h), revealed that 85% of all birds were able to reach the northern fringe of the desert with an intermittent migration strategy. Furthermore, birds stopping over in an oasis (Ouadane, 370 km to the southwest of B?r Amrane) did not carry consistently lower fuel loads compared to the migrants captured in the desert.     

Global navigation in migratory birds: tracks, strategies, and interactions between mechanisms

The advancing development of tracking techniques has led to fascinating new insights into avian migration, documenting the immense diversity, complexity, and flexibility of this phenomenon. Tracking studies so far have confirmed many findings from ringing recoveries and cage studies, for example, the change from flying innate compass courses in the first migration to true navigation, as experienced migrants head toward familiar goals. First attempts to analyze the navigational mechanisms by tracking manipulated migrants indicate strong parallels to those of homing pigeons. Findings suggesting that the magnetic compass of migrants is regularly calibrated by the pattern of polarized light could not be replicated with a number of other birds, pointing out differences between species and possibly region and phases of migration. Tracking has become a valuable tool, complimenting traditional methods by documenting migration behavior in the wild; whether it can be used to further unveil the navigational mechanisms of migrants and the factor used remains an open question.     

Do Seabirds Differ from Other Migrants in Their Travel Arrangements? On Route Strategies of Cory’s Shearwater during Its Trans-Equatorial Journey

Long-distance migrants have developed diverse strategies to deal with the challenges imposed by their annual journeys. These are relatively well studied in some avian groups, such as passerines, shorebirds and raptors. In contrast, few studies have addressed the migratory behaviour of pelagic birds in the light of current theories of optimal migration. Using a dataset of 100 complete migratory tracks gathered along four years, we performed a detailed study on the migratory strategy of a pelagic trans-equatorial migrant, the Cory’s shearwater Calonectris diomedea. We analysed daily routines, stopover ecology and travel speed, as well as the influence of the moon on several behavioural patterns. Cory’s shearwaters adopted a “fly-and-forage” strategy when migrating, similarly to what has been observed in some raptors. However, by flying by dynamic soaring, shearwaters attained high overall migration speeds, and were able to travel thousands of kilometres without making major stopovers and, apparently, without a noticeable pre-migratory fattening period. Other major findings of this study include the ability to adapt daily schedules when crossing major ecological barriers, and the constant adjustment of migration speed implying higher rates of travel in the pre-breeding movement, with a final sprint to the nesting colony. The present study also highlights a preference of Cory’s shearwaters for starting travel at twilight and documents a strong relationship between their migratory activity and the moon phase.