Migratory stopover conditions affect the developmental state of male gonads in garden warblers (Sylvia borin)

Long-distance migrants face the challenge of a short window for reproduction that requires optimal timing and full functional gonads. Male garden warblers (Sylvia borin) meet these demands by initiating testicular recrudescence during spring migration, enabling them to reproduce immediately after arrival at the breeding grounds. In a combined field and laboratory study, we investigated testicular size, plasma levels of luteinizing hormone (LH), androstenedione (AE), 5α-dihydrotestosterone (DHT), testosterone and nocturnal migratory restlessness (Zugunruhe) under different stopover conditions. We manipulated food availability, the duration of stopover and simulated migration by food deprivation. Garden warblers showed significantly retarded testicular development after nine days of stopover under limited food conditions compared to birds that had ad libitum access to food. However, there was no significant difference in Zugunruhe between the two groups. Thus, the degree of Zugunruhe was unaffected by the quality of the stopover site and migration continued independent of the developmental state of the testis. We suggest that male garden warblers face the necessity to either compensate for slowed testicular recrudescence during the subsequent leg of migration and delay arrival at the breeding grounds, or arrive with less developed testes. Either of these may reduce annual reproductive success.     

Flexible remodeling of organ size during spring migration of the garden warbler (Sylvia borin)

The energetic demands of long-distance migratory birds change drastically, depending on the stage of their life cycle. Changing demands are reflected in the up and down regulation of adipose tissue and organ mass. This paper presents new data on organ size changes during different stages of spring migration of garden warblers (Sylvia borin). Phenotypic mass changes were quantified in 13 organs of birds caught in Tanzania, Ethiopia and Egypt. We also sampled birds after a simulated stopover in Egypt. Some organs increased in mass up to about 1.5-fold during migration from Tanzania to Ethiopia, while some remained unchanged or even decreased in mass. During flight across the Sahara, nearly all organ masses including heart and flight muscles were reduced. Exceptionally large reductions (approximately 50%) were observed for liver, bile, spleen, kidney and digestive tract organs. The only exceptions were the testes, which increased 4-fold in mass. During the simulated stopover in Egypt, a significant recovery was observed for kidney, liver, heart, proventriculus, and small intestine. The testes continued to increase in mass. Flexible remodeling of organ size in the course of spring migration thus comprises significant changes for all quantified organs, with a variety of organ-specific patterns. Individual organ patterns are differentially shaped by functional aspects according to the different organ requirements in the alternation of flight and stopover phases, energetics, future demands, and protein requirements. Anticipatory mechanisms account for the size change of the testes, and we suggest the same for the kidney and the gall bladder.     

Basal and stress-induced corticosterone levels of garden warblers,Sylvia borin,during migration

Summary Plasma levels of the metabolically and behaviorally active corticosteroid hormone, corticosterone, were studied in garden warblers in the laboratory and in the field during the autumnal migratory phase. Garden warblers showing nocturnal migratory activity in the laboratory had elevated levels of corticosterone at the end of the dark phase and low levels during daytime. When nocturnal migratory activity was experimentally disrupted by food deprivation and subsequent refeeding or after spontaneous termination of migratory activity this rhythm was absent. Garden warblers stopping over in the Sahara desert during autumnal migration had low levels of corticosterone. Levels were negatively correlated with fat stores and body mass in birds sampled throughout the day. These levels were generally lower than those associated with stress in response to repeated handling and blood sampling. The results suggest (1) the existence of diel changes in adrenocortical hormonal activity that could be involved in regulation of migration, and (2) that garden warblers carrying large fat depots are not stressed by prolonged flight or lack of appropiate feeding areas during migration over the desert.Abbreviations ACTH adrenocorticotrope hormone - cpm counts per minute - EtOH ethanol - RIA radioimmunoassay     

Transition between moult and migration in a long-distance migratory passerine: organ flexibility in the African wintering area

Phenotypic flexibility of organs in migratory birds has been documented for a variety of species of different genera during the migratory period. However, very little is known about phenotypic mass changes of organs with respect to other events within the annual cycle. This seems particularly interesting when birds face different physiological challenges in quick succession. We investigated mass changes of 13 organs from garden warblers (Sylvia borin) during the transition from moult to migration. These long-distance migratory birds perform a complete moult within their wintering area just shortly before the onset of spring migration. Birds were sampled in three successive stages according to their moult status: group I consisted of birds with growing primary or secondary wing feathers, group II consisted of birds with completed wing moult but with still moulting body feathers, and group III consisted of birds that had completed wing moult and body moult. Size-corrected flight muscle, kidney mass, and pancreas mass differed significantly among the three groups. Flight muscle was heaviest in birds that were about to leave their wintering area (group III) compared with birds still in body moult (group II). Kidney and pancreas showed a pattern similar to each other, with the heaviest mass occurring in birds with moulting wing feathers (group I) and significantly reduced mass in birds that had completed wing moult (group II) or both wing and body moult (group III). Mass reductions of kidney and pancreas during the transition from moult to migration are considered to be related to the demands of moult, while increased flight muscle may be due to moult, migration, or both. Phenotypic mass changes of organs in birds occur during their migration, but they also occur during the transition between other phases of the annual cycle such as moult and migration and are not restricted to the flight muscle.     

Circannual variation in blood parasitism in a sub‐Saharan migrant passerine bird,the garden warbler

Knowing the natural dynamics of pathogens in migratory birds is important, for example, to understand the factors that influence the transport of pathogens to and their transmission in new geographical areas, whereas the transmission of other pathogens might be restricted to a specific area. We studied haemosporidian blood parasites of the genera Plasmodium, Haemoproteus and Leucocytozoon in a migratory bird, the garden warbler Sylvia borin. Birds were sampled in spring, summer and early autumn at breeding grounds in Sweden, on migration at Capri, Italy and on arrival and departure from wintering staging areas in West Africa: mapping recoveries of garden warblers ringed in Fennoscandia and Capri showed that these sites are most probably on the migratory flyway of garden warblers breeding at Kvismaren. Overall, haemosporidian prevalence was 39%, involving 24 different parasite lineages. Prevalence varied significantly over the migratory cycle, with relatively high prevalence of blood parasites in the population on breeding grounds and at the onset of autumn migration, followed by marked declines in prevalence during migration both on spring and autumn passage. Importantly, we found that when examining circannual variation in the different lineages, significantly different prevalence profiles emerged both between and within genera. Our results suggest that differences in prevalence profiles are the result of either different parasite transmission strategies or coevolution between the host and the various parasite lineages. When separating parasites into common vs. rare lineages, we found that two peaks in the prevalence of rare parasites occur; on arrival at Swedish breeding grounds, and after the wintering period in Africa. Our results stress the importance of appropriate taxonomic resolution when examining host‐parasite interactions, as variation in prevalence both between and within parasite genera can show markedly different patterns.     

Inability to regain normal body mass despite extensive refuelling in great reed warblers following the trans‐Sahara crossing during spring migration

Migratory birds wintering in Africa face the challenge of passing the Sahara desert with few opportunities to forage. During spring migration birds thus arrive in the Mediterranean area with very low energy reserves after crossing the desert. Since early arrival to the breeding grounds often is of importance to maximize reproductive success, finding stopover sites with good refuelling possibilities after the Saharan passage is of utmost importance. Here we report on extensive fuelling in the great reed warbler Acrocephalus arundinaceus on the south coast of Crete in spring, the first land that they encounter after crossing the Sahara desert and the Mediterranean Sea in this area. Birds were studied at a river mouth and due to an exceptional high recapture rate (45 and 51% in two successive years), we were able to get information about stopover behaviour in 56 individual great reed warblers during two spring seasons. The large proportion of trapped great reed warbler compared to other species and the large number of recaptures suggest that great reed warblers actively choose this area for stopover. They stayed on average 3–4 d, increased on average about 3.5 g in body mass and the average rate of body mass increase was 4.8% of lean body mass d^–1. Wing length affected the rate of increase and indicated that females have a slower increase than males. The results found show that great reed warblers at this site regularly deposit larger fuel loads than needed for one continued flight stage. The low body mass found in great reed warblers (also in birds with high fat scores) is a strong indication that birds staging at Anapodaris still had not been able to rebuild their structural tissue after the strenuous Sahara crossing, suggesting that rebuilding structural tissue may take longer time than previously thought.     

Digestive tract function in the long-distance migratory garden warbler,Sylvia borin

Digestive tract morphology and function of captive garden warblers (Sylvia borin) were measured during four stages of their endogenous circannual rhythm: before, during and after their autumn fattening prior to migration to wintering grounds in Africa, and after a partially simulated migratory flight. Food intake increased by 33% during fattening, utilization efficiency of dry matter tended to increase, and that of energy increased significantly (P0.01). This was because digestive tract capacity (measured as dry tissue mass) increased, so that mean retention time of food remained constant before, during and after fattening (80–84 min). After a 48-h period of starvation of fattened birds to partially simulate a migratory flight, food intake was lower on the first day of refeeding than on the next 4 days, and utilization efficiency was higher on that day, at least partly because of a longer mean retention time (111 min versus 78 min on the third day). Digestive tract dry tissue mass fell by 50% during starvation, and that of the small intestine by 63%. It is concluded that the garden warbler adapts to long-distance migration without feeding by rapidly reducing the size of its digestive tract, an expensive tissue to maintain, during migration in order to save weight and energy, and possibly also to supply part of the fuel and protein required for the flight. The cost of this strategy appears to be the time taken to rebuild the gut at stopover sites with food, but the low probability of finding such a site in the Sahara Desert means that this strategy is probably optimal for garden warblers.     

Annual Cycle and Migration Strategies of a Trans-Saharan Migratory Songbird: A Geolocator Study in the Great Reed Warbler

Recent technological advancements now allow us to obtain geographical position data for a wide range of animal movements. Here we used light-level geolocators to study the annual migration cycle in great reed warblers (Acrocephalus arundinaceus), a passerine bird breeding in Eurasia and wintering in sub-Saharan Africa. We were specifically interested in seasonal strategies in routes and schedules of migration. We found that the great reed warblers (all males, no females were included) migrated from the Swedish breeding site in early August. After spending up to three weeks at scattered stopover sites in central to south-eastern Europe, they resumed migration and crossed the Mediterranean Sea and Sahara Desert without lengthy stopovers. They then spread out over a large overwintering area and each bird utilised two (or even three) main wintering sites that were spatially separated by a distinct mid-winter movement. Spring migration initiation date differed widely between individuals (1-27 April). Several males took a more westerly route over the Sahara in spring than in autumn, and in general there were fewer long-distance travels and more frequent shorter stopovers, including one in northern Africa, in spring. The shorter stopovers made spring migration on average faster than autumn migration. There was a strong correlation between the spring departure dates from wintering sites and the arrival dates at the breeding ground. All males had a high migration speed in spring despite large variation in departure dates, indicating a time-minimization strategy to achieve an early arrival at the breeding site; the latter being decisive for high reproductive success in great reed warblers. Our results have important implications for the understanding of long-distance migrants’ ability to predict conditions at distant breeding sites and adapt to rapid environmental change.     

Environmental regulation of the reproductive system in a flexibly breeding Sonoran Desert bird, the Rufous-winged Sparrow, Aimophila carpalis

We investigated reproductive regulation in male Rufous-winged Sparrows, Aimophila carpalis, a Sonoran Desert passerine that breeds after irregular summer rains. Field and captive data demonstrate that increased photoperiod stimulates testicular development in March and maintains it until early September. Free-living birds caught in July and placed on captive long days (16L: 8D) maintained developed testes for up to 7 months, and free-living birds caught in September, during testicular regression, redeveloped testes when placed on captive long days, indicating that these birds were still photosensitive. Captive birds on long days maintained testicular development when exposed to temperatures mimicking those occurring during regression in free-living birds. In free-living birds, testicular development was observed during spring and summer, but unless this was associated with rainfall, breeding (indicated by juveniles) did not occur. Large increases in plasma luteinizing hormone (LH) in free-living males were correlated with heavy rainfall in July/August, when the birds bred, and in November, when they did not breed. In captive birds, plasma LH concentrations were unresponsive to photoperiodic changes, but may have responded to social cues. Plasma prolactin concentrations were directly correlated with photoperiod in free-living birds, but an effect of photoperiod on prolactin secretion was not seen in captive birds. It is concluded that male Rufous-winged Sparrows use long photoperiods to stimulate and maintain testicular development, but exposure to long photoperiods does not terminate breeding by inducing absolute photorefractoriness. The specific timing of reproductive behaviors is apparently determined by elevated plasma LH coinciding with long day stimulated gonad development.     

Body fat influences departure from stopover sites in migratory birds: evidence from whole-island telemetry

Migration remains one of the great mysteries of animal life. Small migratory birds rely on refuelling stopovers after crossing ecological barriers such as deserts or seas. Previous studies have suggested that fuel reserves may determine stopover duration but this hypothesis could not be tested because of methodological limitations. Here, we provide evidence that subcutaneous fat stores determine stopover duration by measuring the permanence of migratory garden warblers (Sylvia borin) on a small Mediterranean island during spring migration with telemetry methods. Garden warblers with large amounts of fat stores departed the island significantly sooner than lean birds. All except one fat bird left the island on the same evening after capture, with a mean total stopover estimate of 8.8 hours. In contrast, the mean estimated total stopover duration of lean birds was 41.3 hours. To our knowledge, this is the first study that measures the true minimum stopover duration of a songbird during migration.     

Warm Springs,Early Lay Dates,and Double Brooding in a North American Migratory Songbird,the Black-Throated Blue Warbler

Numerous studies have correlated the advancement of lay date in birds with warming climate trends, yet the fitness effects associated with this phenological response have been examined in only a small number of species. Most of these species–primarily insectivorous cavity nesters in Europe–exhibit fitness declines associated with increasing asynchrony with prey. Here, we use 25 years of demographic data, collected from 1986 to 2010, to examine the effects of spring temperature on breeding initiation date, double brooding, and annual fecundity in a Nearctic - Neotropical migratory songbird, the black-throated blue warbler (Setophaga caerulescens). Data were collected from birds breeding at the Hubbard Brook Experimental Forest, New Hampshire, USA, where long-term trends toward warmer springs have been recorded. We found that black-throated blue warblers initiated breeding earlier in warmer springs, that early breeders were more likely to attempt a second brood than those starting later in the season, and that double brooding and lay date were linked to higher annual fecundity. Accordingly, we found selection favored earlier breeding in most years. However, in contrast to studies of several other long-distance migratory species in Europe, this selection pressure was not stronger in warmer springs, indicating that these warblers were able to adjust mean lay date appropriately to substantial inter-annual variation in spring temperature. Our results suggest that this North American migratory songbird might not experience the same fecundity declines as songbirds that are unable to adjust their timing of breeding in pace with spring temperatures.     

The influence of climate on the timing and rate of spring bird migration

Ecological processes are changing in response to climatic warming. Birds, in particular, have been documented to arrive and breed earlier in spring and this has been attributed to elevated spring temperatures. It is not clear, however, how long-distance migratory birds that overwinter thousands of kilometers to the south in the tropics cue into changes in temperature or plant phenology on northern breeding areas. We explored the relationships between the timing and rate of spring migration of long-distance migratory birds, and variables such as temperature, the North Atlantic Oscillation (NAO) and plant phenology, using mist net capture data from three ringing stations in North America over a 40-year period. Mean April/May temperatures in eastern North America varied over a 5°C range, but with no significant trend during this period. Similarly, we found few significant trends toward earlier median capture dates of birds. Median capture dates were not related to the NAO, but were inversely correlated to spring temperatures for almost all species. For every 1°C increase in spring temperature, median capture dates of migratory birds averaged, across species, one day earlier. Lilac (Syringa vulgaris) budburst, however, averaged 3 days earlier for every 1°C increase in spring temperature, suggesting that the impact of temperature on plant phenology is three times greater than on bird phenology. To address whether migratory birds adjust their rate of northward migration to changes in temperature, we compared median capture dates for 15 species between a ringing station on the Gulf Coast of Louisiana in the southern USA with two stations approximately 2,500 km to the north. The interval between median capture dates in Louisiana and at the other two ringing stations was inversely correlated with temperature, with an average interval of 22 days, that decreased by 0.8 days per 1°C increase in temperature. Our results suggest that, although the onset of migration may be determined endogenously, the timing of migration is flexible and can be adjusted in response to variation in weather and/or phenology along migration routes.     

Increased Endoparasite Infection in Late-Arriving Individuals of a Trans-Saharan Passerine Migrant Bird

Earlier migration in males than in females is the commonest pattern in migrating passerines and is positively related to size dimorphism and dichromatism. The early arrival of males is a costly trait that may confer reproductive advantages in terms of better territories and/or mates. Given the physiological cost of migration, early migrants are those in best condition and accordingly the prevalence, load, and/or diversity of parasites is expected to increase in both sexes for late migrants. To test this hypothesis, we sampled 187 trans-Saharan migrant garden warblers Sylvia borin and 64 resident serins Serinus serinus (as a control for potential circannual patterns in parasite load) during spring migration in Spain. We assessed the prevalence of blood parasites (Haemoproteus, Plasmodium, and Leucocytozoon) and the prevalence and load of intestinal parasites (mainly coccidians and spirurids). The relationship between parasite (prevalence, load, and richness) and the timing of passage through a stopover area was tested using generalized linear models. Protandry occurs in the monomorphic garden warbler and males migrated on average 5.5 days before females. Intestinal parasite richness increased with the date of migration. The timing of migration was unrelated to the presence or load of the other parasite groups analyzed. Our results support the idea that the timing of migration is a condition-dependent trait and suggests that multiple intestinal parasite infestations could delay migration in birds. Even in monomorphic species parasites may play a role in sexual selection by delaying the arrival of the most infected individuals at breeding grounds, thereby further increasing the benefits of mating with early-arriving individuals.     

Melatonin reduces migratory restlessness in <Emphasis Type="Italic">Sylvia</Emphasis> warblers during autumnal migration

Introduction

A remarkable aspect of bird migration is its nocturnality, particularly common in Passeriformes. The switch in activity from purely diurnal to also nocturnal is evident even in caged birds that during migratory periods develop an intense nocturnal restlessness, termed Zugunruhe. The mechanisms that control this major change in activity are mostly unknown. Previous work with Sylvia warblers suggested an involvement of melatonin, a hormone associated with day-night cycles in most vertebrates. In a recent study we found no effects of melatonin administration on Zugunruhe during spring migration. However, previous studies indicated that the response to melatonin manipulation could differ between spring and autumn migration, which are in fact separate life history stages. Here we tested whether a non-invasive treatment with melatonin can alter Zugunruhe in wild garden warblers S. borin and blackcaps S. atricapilla subject to temporary captivity at an autumnal stopover site. Food availability in the cage (yes/no) was added as a second factor because previous work showed that it enhanced Zugunruhe.

Results

The melatonin treatment significantly decreased the amount of Zugunruhe, while the availability of food only tended to increase the amount of Zugunruhe. Fuel deposits also had a strong effect on the amount of nocturnal activity: lean birds with a fat score of 1 showed significantly less Zugunruhe than fatter birds. The change in body mass during the time spent in the recording cage depended on food availability, but not on any of the other factors.

Conclusions

This study shows that the migratory programme of two Sylvia warblers can be manipulated by administration of exogenous melatonin and confirms that this hormone is involved in the control of migratory behaviour. To our knowledge, this is one of the first demonstrations that the autumn migratory programme can be altered by hormonal manipulation in migrating birds. The comparison with a similar study carried out with the same modalities during spring migration suggests that there are seasonal differences in the sensitivity of the migratory programme to hormonal factors. In birds breeding in the northern hemisphere, the importance of a timely arrival to the breeding sites could explain why the control of the migratory programme is more rigid in spring.

     

Breeding latitude and timing of spring migration in songbirds crossing the Gulf of Mexico

Each spring, millions of songbirds migrate across the Gulf of Mexico on their way to breeding sites in North America. Data from radar and migration monitoring stations have revealed broad patterns in the spatial and temporal course of trans-Gulf migration. Unfortunately, we have limited information on where these birds have previously spent the winter and where they are migrating to breed. Here we measure stable-hydrogen isotopes in feathers (δD_f) to infer the breeding latitude of five species of songbirds – hooded warblers Wilsonia citrina , American redstarts Setophaga ruticilla , black-and-white warblers Mniotilta varia , ovenbirds Seiurus aurocapilla , and northern waterthrushes S. noveboracensis – that were captured at a stopover site along the coast of southwestern Louisiana in spring 2004. Values of δD_f across all species ranged from −163 to −35‰ (n=212), and within most species the range was consistent with the latitudinal extent of known breeding sites in central and eastern North America. Individuals that arrived first along the northern Gulf coast had δD_f values indicative of southerly breeding sites in hooded warblers, American redstarts, black-and-white warblers, and ovenbirds, but no relationship was found between passage timing and δD_f for northern waterthrushes. Our findings suggest that spring passage is often timed to coincide with the emergence of suitable conditions on breeding areas, with southern breeding birds migrating first.     

Blood parasites in migrating wood‐warblers (Parulidae): effects on refueling,energetic condition,and migration timing

Blood parasites can negatively affect energetic condition, arrival date, and reproductive performance of breeding birds, yet their impact on migrating birds is poorly understood. We quantified haematozoan infection and examined its relationship to migration timing, energetic condition, and refueling rate (plasma triglyceride concentration) for three species of wood‐warblers (Parulidae) migrating through northwestern Ohio, USA from 11 April–3 June of 2004 and 2005. Prevalence of infection for magnolia Dendroica magnolia (36.3%; n=383), yellow‐rumped D. coronata (58.3%; n=518), and yellow warblers D. petechia (12.7%; n=205) was approximately two times higher than reported in other migration studies and similar to infection rates found in breeding areas. Probability and intensity of infection were higher for later migrating yellow‐rumped warblers, especially in younger (second‐year) birds. Furthermore, we found a negative trend between intensity of infection and energetic condition for second‐year yellow‐rumped warblers. Haematozoan infection was not associated with refueling rate for any species. Our results indicate that for some migrants, especially younger birds, probability and intensity of haematozoan infection are related to later migration timing and reduced energetic condition during spring migration, and thus may be important determinants of subsequent reproductive performance in breeding areas.     

The seasonal development of photoperiodic responsiveness in an equatorial migrant, the garden warblerSylvia borin

Summary Garden warblers (Sylvia borin) are migrating European passerines that spend the winter in tropical Africa. To investigate how photoperiod controls the annual cycles of reproduction, molt and migration, garden warblers were exposed to photoperiodic changes simulating those experienced by conspecifics with an equatorial wintering area. At three different times groups of birds were moved from a constant equatorial photoperiod of 12.8 h to a 15-h photoperiod: group 1 on November 19, group 2 on February 24, and group 3 on April 12. In all birds the changes in body weight, nocturnal migratory restlessness, testicular width or diameter of the largest oocyte, and plasma LH concentrations were determined. In addition, the birds were examined for molt. In all birds of groups 2 and 3 the transfer to long photoperiods induced a gonadal cycle associated with spring migratory restlessness and, in the group 3 birds, an increase in LH. In group 1 only few birds showed marginal gonadal responses and the gonadal cycles of the group 2 birds were less pronounced than those of group 3. The results suggest that the hypothalamo-pituitary-gonadal axis was refractory to stimulation by long photoperiods in autumn but, later in the season, became responsive to the same long photoperiods. Unlike the situation in other temperate zone birds, the transition from the photorefractory to the photosensitive state seems to be a gradual process which develops spontaneously and with the appropriate temporal characteristics in the absence of photoperiodic change.Abbreviation LH luteinizing hormone     

A flyway perspective on food resource abundance in a long-distance migrant, the Eurasian teal (Anas crecca)

Two frequent assumptions about the evolution of long-distance migration in birds are that they travel long distances annually to reach food-rich areas for breeding, and that they time their migratory journey to be at staging sites when the latter provide the best feeding conditions. These assumptions have rarely been properly tested, and there is no study in which a species’ major food types have been measured by standardized methods throughout a flyway and over a large part of the year. We here present such data for Eurasian teal (Anas crecca), converted to a common energetic currency, and collected at wintering, spring staging and breeding sites. Teal did not time migration to maximize local food abundance; most birds left wintering and spring staging sites before a sharp increase in invertebrate food abundance occurred. On the other hand, hatching of ducklings coincided with a peak in invertebrate food abundance on boreal breeding lakes. Mean overall food abundance (invertebrates and seeds combined) did not differ between wintering sites in southern France and breeding sites in northern Sweden at the time of breeding. Our results are inconsistent with the hypothesis that long-distance migration in dabbling ducks has evolved because adult birds gain an immediate pay-off in increased food abundance by flying north in spring. However, our data confirm a selective advantage for breeding at higher latitudes, because hatching of ducklings may coincide with a peak in invertebrate emergence and because longer days may increase the duration of efficient foraging.     

A long-distance avian migrant compensates for longitudinal displacement during spring migration

In order to perform true bicoordinate navigation, migratory birds need to be able to determine geographic latitude and longitude. The determination of latitude is relatively easy from either stellar or magnetic cues [1-3], but the determination of longitude seems challenging [4, 5]. It has therefore been suggested that migrating birds are unable to perform bicoordinate navigation and that they probably only determine latitude during their return migration [5]. However, proper testing of this hypothesis requires displacement experiments with night-migratory songbirds in spring that have not been performed. We therefore displaced migrating Eurasian reed warblers (Acrocephalus scirpaceus) during spring migration about 1000 km toward the east and found that they were correcting for displacements by shifting their orientation from the northeast at the capture site to the northwest after the displacement. This new direction would lead them to their expected breeding areas. Our results suggest that Eurasian reed warblers are able to determine longitude and perform bicoordinate navigation. This finding is surprising and presents a new intellectual challenge to bird migration researchers, namely, which cues enable birds to determine their east-west position.     

Carry-Over Effects of Nonbreeding Habitat on Start-to-Finish Spring Migration Performance of a Songbird

For migratory animals, conditions during the nonbreeding period may carry-over to influence spring migration performance. Animals in low-quality habitats are predicted to be in poorer condition, show later migration timing, and travel at slower speeds. This can result in subsequent negative effects on fitness. We tested the hypothesis that nonbreeding season body condition and habitat quality carry-over to affect spring migration performance of a long-distance migratory songbird, the Wood Thrush (Hylocichla mustelina). We tracked individual birds between multiple breeding sites in North America and nonbreeding sites in Central America. First, we compared body condition of nonbreeding birds migrating to the same general region of the breeding range with spring migration performance (timing, speed, and duration) obtained from light-level geolocators. Second, we assessed the Normalized Difference Vegetation Index (NDVI) as a proxy for nonbreeding habitat quality, and predicted that birds from wetter habitat or in wetter years (higher NDVI) would show improved migration performance relative to birds from drier sites. We found no evidence of individual-level carry-over effects of nonbreeding season body condition on spring migration performance. Lower NDVI of nonbreeding habitat resulted in delayed spring migration departure, but this effect disappeared by arrival at breeding sites. Birds occupying drier nonbreeding sites migrated faster and for fewer days, compensating for their relatively late departure. We also documented a broader pattern in NDVI and migration timing and distance, in that birds that occupied the wettest areas in the southern part of the nonbreeding range departed significantly later and migrated farther. Our results suggest that individual carry-over effects of nonbreeding habitat quality may be compensated for by a faster and shorter migration strategy. At a broad scale, consistently later spring timing and longer migration distances were associated with the wettest areas (the highest quality habitats) of the Wood Thrush non-breeding range. This supports the theory that high-quality habitats offset the costs of farther migration, resulting in a leap-frog migration pattern.