Energetics of fattening and starvation in the long-distance migratory garden warbler,Sylvia borin,during the migratory phase

Garden warblers (Sylvia borin) were subjected to starvation trials during their autumnal migratory phase in order to simulate a period of non-stop migration. Before, during and after this treatment the energy expenditure, activity, food intake and body mass of the subjects were monitored. Assimilation efficiency was constant throughout the experiments. The catabolized (during starvation) and deposited body tissue (during recovery) consisted of 73% fat. Basal metabolic rate was decreased during the starvation period and tended to a gradual increase during the recovery period. The reduced basal metabolic rate can possibly be attributed to a reduced size/function of the digestive system, which is consistent with the sub-maximal food intake immediately after resuming the supply of food to the experimental birds. The observed reductions in basal metabolic rate during starvation and activity during recovery can be viewed as adaptations contributing to a higher economization of energy supplies. The experimental birds were unable to eat large quantities of food directly after a period of starvation leading to a comparatively low, or no increase in body mass. Such a slow mass increase is in agreement with observations of migratory birds on arrival at stop-over sites.Abbreviations BM body mass - BMR basal metabolic rate - LBM lean body mass - RQ respiratory quotient     

The autumnal fattening of the long-distance migratory garden warbler (Sylvia borin) is stimulated by intermittent fasting

To investigate the proximate influence of a changing food availability on the seasonal fattening of migratory birds, garden warblers (Sylvia borin) following postnuptial moult were food restricted once a week. Body mass, food intake, plasma hormone and metabolite levels were measured and compared to birds which always had ad libitum food access. The food-restricted birds increased their body mass significantly earlier than the controls. The accelerated fattening was initially not accompanied by hyperphagia and may be due to either an increased food utilisation efficiency or a reduced metabolic rate. An increase of basal glucagon and corticosterone and a decrease of insulin levels prior to fattening were not significant, however, they resulted in a significant decrease of the insulin:glucagon ratio. This ratio was also lower in food-restricted birds than in control birds and may account for the difference in the fattening progress. We conclude that seasonal fattening may be stimulated by a catabolic impulse which could be imposed in free-living birds by a decrease of food availability and/or by an increase of energy expenditure. A negative energy balance is hypothesised to be a common proximate factor affecting migratory as well as winter fattening. Accepted: 24 August 2000     

Effects of food-deprivation on migratory restlessness and diurnal activity in the garden warbler Sylvia borin

Summary The effects of tadpole body size, tadpole sibship, and fish body size on predation of gray treefrog tadpoles, Hyla chrysoscelis, were studied in laboratory and artificial pond experiments. Tadpole body size had a significantly positive effect on the survival of tadpoles in all experiments. The relationship between tadpole biomass eaten and biomass available suggested that fish were not satiated when consuming the largest tadpoles. Large tadpoles were probably better able to evade predators. A difference in survival among full sib families of tadpoles was only present in one family, suggesting that genetic differences in predator avoidance behavior or palatability were probably secondarily important to body size per se. Fish body size had a significantly negative effect on the survival of tadpoles. Larger fish consumed a larger number and proportion of tadpoles as well as greater biomass. These results indicate that environmental factors affecting the growth rate of tadpoles cand dramatically alter their vulnerability to gape-limited predators.     

Nutritional requirements for maintenance of body weight and fat deposition in the long-distance migratory garden warbler, Sylvia borin (Boddaert)

Intake of food, protein, fat and carbohydrates and their fecal output and the birds' weights were recorded during different feeding trials with specific nutrient reduced diets in the old-world long-distance migratory garden warbler. The birds' body weights were affected by low dietary protein as well as low dietary fat levels. Low dietary protein and fat levels were associated with significant changes in daily gross and net food intake and in the efficiency of food and nutrient utilization. Birds fed on diets with low nutrient levels for an extended length of time recovered in weight after an initial weight loss. They obviously compensated the restricted nutrient levels primarily by increasing the daily food intake and by changing the efficiency of food and nutrient utilization. Effects of restricted dietary nutrient levels on body weight and adaptation depended on the previous composition of the food. The average daily net fat intake was much higher than the average daily net protein intake, both for maintenance of a constant body weight and for successful regain of weight. The data were further discussed with respect to the role of a fruit diet in omnivorous passerine birds.     

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.     

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.     

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     

When and where to fuel before crossing the Sahara desert – extended stopover and migratory fuelling in first-year garden warblers Sylvia borin

Large numbers of passerine migrants cross the Sahara desert every year on their way to-and-from wintering areas in tropical Africa. In the desert, hardly any fuelling opportunities exist and most migrants have to prepare in advance. A central question is how inexperienced birds know where to fuel. Inexperienced garden warblers Sylvia borin were studied in Greece just before the desert crossing in autumn. Body mass data collected at two sites indicate that most birds do not fuel for the desert crossing further north. For the first time, detailed information about stopover duration close to the Sahara desert was studied by using light weight radio-transmitters. Results from Crete show that most first-year garden warblers arrive with relatively small fuel loads in relation to lean body mass (<30%), stay for 13–20 d and depart with an average fuel load of about 100%. Radio-tagged birds performed small scale movements initially and took advantage of fig fruits. Birds trapped at fig trees were heavier than birds trapped with tape lures, showing that tape lures can bias the sample of migrants trapped. The precise fuelling pattern found indicates that first-year migrants must also include external spatial cues to make the preparation for crossing the desert in the right area.     

Effects of experience and object complexity on exploration in garden warblers (Sylvia borin)

Animals are increasingly exposed to novel environmental conditions because of human activities. An organism's ability to cope with these challenges is critical to its survival. The decision of whether or not an animal should investigate environmental changes involves a trade-off between the risk and the potential benefit of a novel resource. Novelty often elicits approach (exploratory) as well as avoidance behaviour in animals, and the extent of each of these behaviours may influence learning about the environment. We formulated and tested several hypotheses to determine how external factors (complexity of a novel stimulus) and internal factors (experience with objects) might influence exploration latencies to touch a novel object based on either cost or benefit considerations. Garden warblers were confronted with a simple and a complex novel object in their familiar cages, and latencies until first touch were measured. Additionally, corticosterone levels before and after the experiment were measured to assess whether differences in baseline levels might explain variations in latency to approach and whether presentation of novel objects is stressful. The complex object was touched significantly later than the simple object. Experienced birds investigated both types of objects later than inexperienced birds. Although birds showed marked approach–avoidance behaviour with the complex object, none of the objects elicited a corticosterone stress response. The results indicate that garden warblers consider both costs and benefits when exploring novel objects, and that the relative influence of costs and benefits varies with external and internal factors.     

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.     

Differential catabolism of muscle protein in Garden Warblers (Sylvia borin): flight and leg muscle act as a protein source during long-distance migration

Samples of flight and leg muscle tissue were taken from migratory garden warblers at three different stages of migration: (1) pre-flight: when birds face an extended flight phase within the next few days, (2) post-flight: when they have just completed an extended flight phase, and (3) recovery: when they are at the end of a stop-over period following an extended flight phase. The changes in body mass are closely related to the changes in flight (P<0.001) and leg muscle mass (P<0.001), suggesting that the skeletal muscles are involved in the protein metabolism associated with migratory flight. From pre- to post-flight, the flight and the leg muscle masses decrease by about 22%, but are restored to about 12% above the pre-flight masses during the recovery period. Biochemical analyses show that following flight a selective reduction occurred in the myofibrillar (contractile) component of the flight muscle (P<0.01). As this selective reduction accounts only for a minor part of the muscle mass changes, sarcoplasmic (non-contractile) and myofibrillar proteins of both the flight and leg muscle act as a protein source during long-distance migration. As a loss of leg muscle mass is additionally observed besides the loss in flight muscle mass, mass change seems not to be strictly associated with the mechanical power output requirements during flight. Whereas the specific content of sarcoplasmic proteins in the flight muscle is nearly twice as high as that in the leg muscle (P<0.001), the specific content of myofibrillar proteins differs only slightly (P < 0.05), being comparably low in both muscles. The ratio of non-contractile to contractile proteins in the flight muscle is one of the highest observed in muscles of a vertebrate.     

Advanced departure dates in long-distance migratory raptors

Evidences for phenological changes in response to climate change are now numerous. One of the most documented changes has been the advance of spring arrival dates in migratory birds. However, the effects of climate change on subsequent events of the annual cycle remain poorly studied and understood. Moreover, the rare studies on autumn migration have mainly concerned passerines. Here, we investigated whether raptor species have changed their autumn migratory phenology during the past 30 years at one of the most important convergent points of western European migration routes in France, the Organbidexka pass, in the Western Pyrenees. Eight out of the 14 studied raptor species showed significant phenological shifts during 1981–2008. Long-distance migrants displayed stronger phenological responses than short-distance migrants, and advanced their mean passage dates significantly. As only some short-distance migrants were found to delay their autumn migration and as their trends in breeding and migrating numbers were not significantly negative, we were not able to show any possible settling process of raptor populations. Negative trends in numbers of migrating raptors were found to be related to weaker phenological responses. Further studies using data from other migration sites are necessary to investigate eventual changes in migration routes and possible settling process.     

Restricted dispersal in a long-distance migrant bird with patchy distribution,the great reed warbler

In patchily distributed species dispersal connects local populations into metapopulations. Reliable quantifications of dispersal are therefore crucial to understanding the population dynamics and genetic structure of such metapopulation systems. The great reed warbler (Acrocephalus arundinaceus) inhabits eutrophic lakes and has a patchy breeding distribution. In this study we investigated the dispersal pattern of the great reed warbler based on an extensive capture-recapture effort covering a large census area (22,500 km²). At two adjacent breeding sites (10 km apart) in southern Central Sweden, the "main study area", we ringed the majority of adult and nestling great reed warblers between 1992 and 1999. In 1998 and 1999, we opportunistically searched for territorial males at the majority of the Swedish breeding sites, and were able to examine about 56% of all males in the region. Analyses of recaptured males demonstrated that philopatry predominated. Sixty-nine percent of the recruiting nestlings returned to breed in the main study area (their natal area), and 92% of the resighted adults were found at the same breeding locality in both study years. Breeding dispersal was significantly more restricted than natal dispersal. Additional data on natal and breeding dispersal within the main study area in 1992-1999 suggested that females were as philopatric as males. The overall high level of philopatry, with only occasional longer dispersal distances documented, yielded a root-mean-square dispersal distance of 33 km per generation. High philopatry, short dispersal distances and similar dispersal patterns of male and female great reed warblers contrast the findings among birds in general, but conform to data of species having patchy breeding habitat and isolated populations. Restricted dispersal suggests limited gene flow even among several Swedish populations, which is in line with some previous findings of the population ecology of the great reed warbler: (1) structured mtDNA lineages among European populations; (2) small-scale population differences in song patterns; and (3) low genetic variation and occurrence of inbreeding depression in our main study population.     

Macroecology of habitat choice in long-distance migratory birds

Patterns of habitat choice in ecological communities are not only influenced by present-day selective forces but also by historical processes, such as the biogeographical history of the lineages they are composed of. Nevertheless, it has been very difficult to test historical factors. The possible tropical origin of long-distance migratory birds provides an opportunity for such a test. If habitat choice of long-distance migrants is inherited from their tropical ancestors then Nearctic long-distance migrants might have acquired their habitat choice from Neotropical forest species and European long-distance migrants from African savannah species. Here we use a macroecological approach to show that this hypothesis can be confirmed. Long-distance migrants in the Nearctic are found in forested habitat types, while those in Europe are found in open ones. In comparison, the habitat choice of residents and short-distance migrants (in genera without long-distance migration) does not differ between the Nearctic and Europe. These results demonstrate that habitat choice in temperate bird communities can be explained by the tropical history of long-distance migrants. Thus, habitat choice seems to be shaped not only by local mechanisms, but also by processes acting on much larger spatial and temporal scales.     

Spring migration of the Garden Warbler(Sylvia borin) across the Mediterranean Sea

Summary Information concerning the strategy adopted by passerine migrants in crossing ecological barriers is still scant, especially as far as spring migration is concerned. With the aim of investigating the pattern of desert and sea crossing in spring, capture frequency, biometry and physical conditions of Garden Warblers ringed at 18 stations spread over the Western and Central Mediterranean were analysed as a function of the geographical position of the ringing sites. The patterns of increasing capture frequency and wing length with longitude, matching those recorded within the species' wide breeding range, suggest wide-front movements across the Sahara and Mediterranean. The progressively delayed trapping hour with latitude indicates that Garden Warblers migrate actively during the day. Despite the potentially favourable spring ecological conditions, the North African coast seems not to be used substantially to restore the reserves depleted during the desert crossing. According to recent estimates of flight range, fat levels recorded at our Mediterranean stations are compatible with the absence of refuelling in North Africa, assuming a fat load at departure of about 27 g or lower fat load and some tail wind assistance.

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Frühjahrszug der Gartengrasmücke(Sylvia borin) über das Mittelmeer

Zusammenfassung Noch wie vor ist wenig bekannt, wie Singvögel große ökologische Barrieren überwinden. Dies gilt besonders für den Frühjahrszug. Um zu untersuchen, wie im Frühjahr die Wüste und das Mittelmeer überquert werden, haben wir Fangfrequenz, Biometrie und physische Kondition von Gartengrasmücken analysiert, die an 18 verschiedenen Fangstationen im zentralen und westlichen Mittelmeer gefangen wurden. Mit zunehmender geographischer Länge nahmen die Fanghäufigkeit und die Flügellänge zu. Diese Zunahmen entsprechen denen im Brutgebiet und zeigen, daß Gartengrasmücken die Sahara und das Mittelmeer in breiter Front überqueren. Die mit zunehmender geographischer Breite spätere Tagesstunde des Fangs zeigt, daß Gartengrasmücken auch tags ziehen. Ungeachtet der möglichen günstigen Rastbedingungen im Frühjahr in N-Afrika scheint die nordafrikanische Küste aber nicht zur Rast und Wiederauffüllung verbrauchter Energievorräte genutzt zu werden. Die gefundenen Fettwerte und die damit mögliche Schätzung der potentiellem Flugstrekke deuten eher darauf hin, daß die Gartengrasmücken südlich der Sahara mit einer Körpermasse von etwa 27 g oder weniger starten und unter Nutzung von Rückenwinden über Sahara und Mittelmeer ohne weiteres Auftanken ziehen.

     

Compensatory frugivory in migratory Sylvia warblers: geographical responses to season length

Seasonal shifts in the diets of Garden Warblers Sylvia borin , Blackcaps S. atricapilla and Lesser Whitethroats S. curruca at a North German site were studied from samples obtained by flushing the digestive tract of the birds. The significance of seven species of berries and pulpy fruits in the diet of these species while feeding in riparian vegetation during the premigratory and migratory seasons was assessed and compared with the pattern of frugivory shown by the same species in southern Germany. Because of the short time span between hatching and onset of migration, first-year long- and medium-distance migratory Sylvia warblers from northern Europe are forced to accelerate their juvenile development. Presumably to meet the higher nutritional demands of an accelerated development, juvenile Garden Warblers and Blackcaps in northern Germany ate relatively more fruit during premigration than Blackcaps in southern Germany. Fruit occurred in about 90% of their diet samples during both premigration and migration. However, this might not hold true for experienced adult birds. Garden Warblers accelerated the rate of fattening and reduced the time needed to prepare for migration when feeding on a mixed fruit diet. There were indications that Blackcaps directly influenced the fruit choice of Garden Warblers when fruits preferred by both species were scarce. Blackcaps and Garden Warblers showed subtle differences in their seasonal preferences for black elder Sambucus nigra and woody nightshade Solanum dulcamara .     

Age-related variation in wing shape of migratory and sedentary Blackcaps Sylvia atricapilla

In many passerines, juveniles have shorter and more rounded wings than adults. Given that (1) long and pointed wings improve endurance in migratory flights, (2) shorter and rounded wings improve manoeuvrability, and (3) juvenile birds are more vulnerable to predators than adults, it has been hypothesised that ontogenetic variation in wing shape results from a greater importance of predation avoidance relative to migration performance during the first year of life. If so, wing shape should not change with age in the absence of migration-related selection for longer and more pointed wings. We test this by studying the variation with respect to age in wing length and wing pointedness of migratory and sedentary Blackcaps wintering in southern Spain. Migratory Blackcaps had longer and more pointed wings than sedentary Blackcaps. Juveniles had shorter wings than adults in migratory populations, but not in sedentary populations. The variation with age in wing pointedness was less pronounced, and was found in migratory females only. These differences between the two traits could be related to a stronger selection for pointed wings than for longer wings with increasing distance of migration, and to an increased migratoriness of females in partially migratory Blackcap populations. We hypothesise that, in migratory Blackcaps, a shorter and more rounded wing in juveniles could be selected for if the decrease in predation rate compensated for the somewhat greater costs of the first migration attempt. On the other hand, there are no costs of migration in sedentary Blackcaps, which hence maintain a similar wing shape, giving high manoeuvrability, both as juveniles and as adults.