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.     

Wind conditions experienced during the day predict nocturnal restlessness in a migratory songbird

A variety of methods have been used to study the relationship between wind conditions and departure decisions of migrant birds at stopover sites. These methods are either costly or suffer from inaccuracy in determining whether or not an individual has resumed migration. Here we present a novel and simple approach to studying the relationship between wind conditions and departure likelihood. Northern Wheatears Oenanthe oenanthe caught during stopover were temporarily caged to measure their nocturnal migratory restlessness, which is an accurate proxy for their individual departure likelihood. We then related the degree of nocturnal restlessness to wind conditions prevailing at the time of capture. Confirming the general pattern from previous studies of departure, the intensity of nocturnal migratory restlessness, and hence departure likelihood, increased with increasing wind support towards the migratory goal. This suggests that approximating the propensity to depart by measuring nocturnal migratory restlessness is a reliable way to study the effect that wind conditions experienced during stopover has on the departure decision of migrants. Our study also shows that nocturnal migrants possess the ability to use information gathered during the day for their departure decisions at night. Because measuring migratory restlessness is straightforward and inexpensive, our approach is ideally suited to test hypotheses regarding spatio‐temporal variation in wind selectivity in migrating birds.     

Geomagnetic information modulates nocturnal migratory restlessness but not fueling in a long distance migratory songbird

Geomagnetic cues have been shown to influence migratory orientation and migratory fuelling in night‐migratory songbird species. Here, we used captive‐bred northern wheatears Oenanthe oenanthe from the southern Norwegian population to show that other aspects of the birds’ migratory program can be influenced by magnetic cues as well. We observed that the amount of migratory restlessness increased strongly with progression of the migratory season when the birds were kept constantly in the magnetic field of northern Germany, but the amount of migratory restlessness decreased when the magnetic field changed along the birds’ natural flyway are simulated. Thus, the Earth's magnetic field can also act as a ‘signpost’ cue for fine‐tuning the spatio‐temporal course of migration.     

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.     

Pinealectomy abolishes the circadian rhythm of migratory restlessness

Summary Pinealectomy of White-throated Sparrows (Zonotrichia albicollis) free-running under constant conditions in dim light abolishes the circadian rhythm of nocturnal spring and fall migratory restlessness (Zugunruhe) as well as the rhythm of summer daytime locomotor activity (Pigs. 1 and 2). Rhythmicity persists in sham-operated birds. Pinealectomized birds are synchronized by a light cycle but their activity rhythm decays to arrhythmicity when they are released from entrainment into constant dim light. The pineal of the white-throat seems essential for the expression of circadian rhythms of both daytime activity and migratory restlessness. These findings support the hypothesis that the avian pineal is fundamentally involved in circadian organization.     

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     

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.     

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.     

Potential age differences in the migratory behaviour of a nocturnal songbird migrant during autumn and spring

In migratory songbirds, older individuals are thought to be more efficient migrants than younger individuals. Age‐specific differences in migratory efficiency have been reported mainly in respect of arrival timing, energy stores, rate of energy accumulation, departure behaviour, and departure direction. Yet, these traits were rarely assessed simultaneously in a single species. We further lack information whether age‐specific differences in behavioural traits present in autumn still manifest to the same degree in spring. Here we used the northern wheatear Oenanthe oenanthe, a long‐distance nocturnal songbird migrant, and investigated age‐specific differences in energy stores at capture (autumn: 1059 birds/spring: 803 birds), rate of energy accumulation (168/147), nocturnal departure timing (126/105), and departure direction (94/77) for both seasons. We found that in autumn, young northern wheatears departed significantly later in the night than older birds. This difference was not observed in spring. The resulting advance in nocturnal departure timing by young birds from autumn to spring may be due to learning based on experience gained during autumn, and/or to selective disappearance of those young individuals showing late departure times during autumn. We found no age‐specific difference in any of the other migratory traits investigated. To get a better understanding of age effects in songbird migrants, we need to study the potential adjustments of migratory behaviour within the individual over its life time. By individually tracking songbirds throughout their lifetime, we could start estimating whether a certain migratory decision (fuelling, departure, orientation) translates into higher (or lower) fitness, and whether individuals adjust their migratory behaviour based on learning from ‘wrong decisions’.     

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.     

Heat loss in sleeping garden warblers (Sylvia borin) during migration

For small songbirds, energy is often a limiting factor during migration and, for this reason, they are forced to alternate nocturnal flights with stopovers to rest and replenish energy stores. Stopover duration has a key role for a successful migration and may have an important impact on fitness. Thus, migrants need to optimize their energy consumption at this stage to reduce their permanence at the site. A recent study has shown that lean individuals reduce their metabolic rate when tucking the head in the feathers during sleep. The underlying mechanism is very likely a reduction in conductance, but the thermoregulatory benefit of the increased insulation has never been quantified yet. Here, we compared heat loss in individual migratory birds while sleeping in different postures. Using a thermal camera and a within-individual approach, we estimated that Garden Warblers can reduce their rate of heat loss by 54% by sleeping with the head tucked in the feathers. This energy saving has a relevant impact on the individual’s energy balance because it can account for up to 8.69% of daily energy expenditure during stopover. Our study provides novel and important information to understand the fundamental role of thermoregulatory strategies on bird’s energy management.     

Body condition and wind support initiate the shift of migratory direction and timing of nocturnal departure in a songbird

1.?An innate migration strategy guides birds through space and time. Environmental variation further modulates individual behaviour within a genetically determined frame. In particular, ecological barriers could influence departure direction and its timing. A shift in the migratory direction in response to an ecological barrier could reveal how birds adjust their individual trajectories to environmental cues and body condition. 2.?Northern wheatears of the Greenland/Iceland subspecies Oenanthe oenanthe leucorhoa arrive in Western Europe en route from their West African winter range. They then undergo an endogenously controlled shift in migratory direction from north to north-west to cross a large ecological barrier, the North Atlantic. We radiotracked these songbirds departing from Helgoland, a small island in the North Sea, over an unprecedented range of their journey. 3.?Here, we show that both birds' body condition and the wind conditions that they encountered influenced the departure direction significantly. Jointly high fuel loads and favourable wind conditions enabled migrants to cross large stretches of sea. Birds in good condition departed early in the night heading to the sea towards their breeding areas, while birds with low fuel loads and/or flying in poor weather conditions departed in directions leading towards nearby mainland areas during the entire night. These areas could be reached even after setting off late at night. 4.?Behavioural adjustment of migratory patterns is a critical adaptation for crossing ecological barriers. The observed variation in departure direction and time in relation to fuel load and wind revealed that these birds have an innate ability to respond by jointly incorporating internal information (body condition) and external information (wind support).     

Seasonal differences in energy requirements of Garden Warblers Sylvia borin migrating across the Sahara desert

The Sahara desert acts as an ecological barrier for billions of passerine birds on their way to and from their African wintering areas. The Garden Warbler Sylvia borin is one of the most common migrants involved. We used body mass of this species from Greece in autumn and spring to simulate the desert crossing and to assess how body mass relates to fuel requirement. The flight range estimates were adjusted to the seasonal extent of the desert, 2200 km in autumn and about 2800 km in spring. In autumn, with an average fuel load of about 100% of body mass without fuel, birds were not able to cross the desert in still air, but northerly winds prevail during September and with the average wind assistance only one in 14 was predicted to fail to make the crossing. Body mass data from spring, after the desert crossing, was used to estimate departure body mass from south of the desert. The average wind assistance in spring is close to zero and departure body mass of the average bird arriving at Antikythira, a small Greek island, under such conditions was estimated to be 34.6 g, which corresponded to a fuel load of 116%. Calculations based on 1% body mass loss per hour of flight showed slightly larger body mass loss than that calculated from flight range estimates. The results suggest that passerine birds about to cross the eastern part of the Sahara desert need to attain a larger fuel load in spring than in autumn.