Effect of the Social Environment on Spring Migration Timing of a Songbird

The influence of the social environment on the timing of the annual cycle is poorly understood. Seasonally migratory birds are under pressure to accurately time their spring migration, and throughout the annual cycle, they may experience variability of the local sex-ratio. A population-level male-biased sex ratio is predicted to advance spring migration timing in males and is attributed to the increased intra-specific competition for access to females and/or breeding territories. The present study had two goals. First, to develop a method that utilizes digitally coded radio-transmitters to quantify the activity of flocked individuals in captivity. Second, to use this method to test the hypothesis that the social environment influences the spring migration traits of male yellow-rumped warblers (Setophaga coronata coronata). To accomplish this, birds were captured in the fall in Long Point, Ontario, and transferred to the Advanced Facility for Avian Research, London, Ontario. In the winter, they were assigned to a slightly male- or female-biased treatment and housed in flocks in large free-flight rooms. Throughout the experimental period, we took body mass measurements and standardized photos to monitor body condition and molt progression. To measure locomotor activity, the birds were outfitted with digitally coded radio-transmitters in April and photo-triggered to enter a migratory phenotype. The tagged birds were released at their capture site in May and the Motus Wildlife Tracking System was used to determine stopover departure timing and migratory movements. Sex ratio did not influence body mass or molt progression. However, males from the male-biased treatment had significantly less locomotor movement than those from the female-biased treatment. Additionally, a lower proportion of males from the male-biased treatment initiated migratory restlessness, an indicator of the urge to migrate. Overall, these findings suggest that the social environment can influence behavior of songbirds, but do not support the hypothesis that a male-biased sex ratio accelerates migration.

Extrinsic factors, endocrine mechanisms, and behavioral indicators of migratory restlessness in wintering whooper swans (Cygnus cygnus)

Extrinsic factors, endocrine mechanisms, and behavioral indicators of migratory restlessness were studied in wintering whooper swans (Cygnus cygnus) in the Sanmenxia Swan National Wetland Park in western Henan Province, central China. First, the fecal glucocorticoid metabolite (FGM) concentration was established and related to mean air temperature or photo period (day length) using simple linear or non-linear regression models. After a model selection procedure, the best fitted model revealed that an increase of FGM concentration was associated with an increase in the squared mean air temperature (R2 = 0.88). Other models showed an increasing FGM concentration to correspond with increasing values of day length, squared day length, and mean air temperature—however without statistical support. In a second step, behavioral frequencies of seven behaviors were condensed into three behavioral principal components (PCs) using principal components analysis. Behavioral PCs largely corresponded to three activity phases described for wintering whooper swans in central China and were correlated with the FGM concentration using Spearman's rank-order correlations. Results revealed a significant correlation between FGM and behavioral PC2 (positive factor loadings from vigilance and preening, negative loading from foraging). Finally, we tested for an effect of behavioral PCs on changes in winter home range size using a set of multiple linear regression models. Results of averaged model parameter estimates showed only the behavioral PC3 (positive factor loadings from fighting and calling, negative loading from locomotion) had a marginal significant effect on home range size. Results confirmed findings of previous studies on migratory restlessness in whooper swans. However, due to the small sample size (N = 15 weeks) the effect of PC3 on home range size was weak and should be viewed with caution.

Flexibility and Control of Circadian Activity, Migratory Restlessness and Fueling in Two Songbird Migrants

Juvenile songbirds rely on an endogenous program, encoding direction, distance, fueling, and timing of migration. Migratory distance is species-specific, expressed as a period of migratory restlessness, for which the length is correlated with distance, while fueling is modified to meet anticipated flight distances controlled by geomagnetic cues and amount of day-light available for foraging. How daylength affect onset and level of migratory activity and fueling decisions in wild birds have so far received limited attention. Here we study how photoperiod controls onset, level and extent of autumn migratory activity and fueling in juvenile diurnally migrating dunnocks, and nocturnally migrating European robins by experimentally increasing daylength. For both species, we kept a control group indoors at the location of capture in southern Sweden exposed to the natural photoperiod, and an experimental group with increased and advanced photoperiod by 2 h in the morning. Dunnocks initiated migratory activity at sunrise (or artificial sunrise) in both groups, demonstrating a highly responsive and flexible component for the onset of migration triggered by light. Experimental robins anticipated the end of nocturnal migratory activity predicting the earlier sunrise immediately after the time-shift and expressed this behavior already under darkness, supporting a fast-resetting mechanism to the new diel period. Timing of end of morning activity was not affected by the earlier sunrise in both species, suggesting a fixed endogenous control that persisted throughout the 13-day study period. Experimental dunnocks expressed higher overall activity and lower fuel loads than controls, while robins did not change their overall activity and fuel load in response to the shifted and increased photoperiod. These results reveal important adaptations for circadian timekeeping including both a flexible onset open to fast modifications and a more rigid end, with differential effects by the treatment on migratory activity and fueling in the two species.

Antibiotics affect migratory restlessness orientation

Magnetoreception is a sense that allows the organism to perceive and act according to different parameters of the magnetic field. This magnetic sense plays a part in many fundamental processes in various living organisms. Much effort was expended in finding the ‘magnetic sensor’ in animals. While some experiments show a role of the ophthalmic nerve in magnetic sensing, others show that effects of light on processes in the retina are involved. According to these inconclusive and puzzling findings, the scientific community has yet to reach an agreement concerning the underlying mechanism behind animal magnetic sensing. Recently, the symbiotic magnetotaxis hypothesis has been forwarded as a mechanistic explanation for the phenomenon of animal magnetoreception. It suggests a symbiotic relationship between magnetotactic bacteria (MTB) and the navigating host. Here we show that in contrast to the control group, antibiotic treatment caused a lack of clear directionality in an Emlen funnel experiment. Accordingly, the antibiotics treatment group showed a significant increase in directional variance. This effect of antibiotics on behaviors associated with animal magnetic sensing is, to the best of our knowledge, the first experimental support of the symbiotic magnetotactic hypothesis.

The amount of available food affects diurnal locomotor activity in migratory songbirds during stopover

Migratory passerine birds fly long distances twice a year alternating nocturnal flights with stopovers to rest and replenish energy stores. The duration of each stopover depends on several factors including internal clocks, meteorological conditions, and environmental factors such as availability of food. Foraging entails energetic costs, and if birds need to refuel efficiently, they should modulate their activity in relation to food availability. We investigated how food availability influences locomotor activity in migrating birds of six passerine species at a spring stopover site in the central Mediterranean Sea. We selected birds with low fat scores which we expected to be strongly motivated to refuel. We simulated stopover sites of different quality by providing temporarily caged birds with different amounts of food to simulate scarce to abundant food. We analysed the diurnal locomotory activity as a proxy for food searching effort. Low food availability resulted in an increased diurnal locomotor activity in almost all species, while all birds showed low intensity of nocturnal migratory restlessness. In conclusion, our study shows that food availability in an important determinant of behaviour of migratory birds at stopover sites.

Variation in chronotype is associated with migratory timing in a songbird

Like many organisms, birds exhibit daily (circadian) and seasonal biological rhythms, and within populations both daily and seasonal timing often vary among individuals. Because photoperiod interacts with the circadian rhythms of many organisms to induce seasonal changes in behaviour and physiology, it is hypothesized that differences in daily timing, called chronotypes, underpin differences among individuals in the timing of seasonal events. For seasonal events stimulated by increasing daylength, this hypothesis predicts a positive relationship between the timing of daily and seasonal activities of individuals, with advanced chronotypes expressing events earlier in the year. The few previous tests of this hypothesis have focused on seasonal reproductive timing in birds. However, the hypothesis predicts that this relationship should extend to other photoinduced seasonal events. Therefore, we tested whether variation in chronotype was associated with variation in spring migratory timing in a captive songbird model, the pine siskin ( Spinus pinus ). We found that pine siskins expressing migratory restlessness exhibited repeatable chronotypes in their timing of nocturnal activity. Further, chronotype was significantly associated with the onset date of migratory behaviour, consistent with the hypothesized relationship between chronotype and seasonal timing.

Sex differences in migratory restlessness behavior in a Nearctic–Neotropical songbird

Most seasonally migrating songbirds exhibit protandry, whereby males arrive to breeding sites in the spring before females. The proximate behavioral mechanisms of protandry are largely unknown for most species, but could include earlier migratory departure from wintering sites by males or overall faster migration by males. Using onset and intensity of migratory restlessness as proxies for departure timing and rate of migration, respectively, we evaluated these 2 hypothesized mechanisms in a Nearctic–Neotropical migrating songbird, the Black-throated Blue Warbler (Setophaga caerulescens). Birds were captured during fall migration, held in captivity over winter, and photostimulated in the spring to induce migratory behavior. Video analysis was used to separately quantify stereotypical nocturnal wing whirring and jumping migratory restlessness behaviors. The birds were then radio-tagged and released in mid-May to compare stopover duration between the sexes and validate migratory restlessness in captivity as a proxy for the motivation to migrate in the field. In captivity males initiated migratory restlessness earlier in the spring than females, demonstrating innate differences in the onset of spring migration in this species. Males also displayed higher-intensity wing whirring behavior, suggesting potential sex differences in flight behavior that could influence migration rate. We found no sex differences in stopover duration in the field following release. However, stopover duration was negatively correlated with total migratory restlessness intensity on the last night the birds were held in captivity, which supports migratory restlessness as a proxy for the motivation to migrate at the individual level.

Magnetic storms disrupt nocturnal migratory activity in songbirds

Birds possess a magnetic sense and rely on the Earth's magnetic field for orientation during migration. However, the geomagnetic field can be altered by solar activity at relative unpredictable intervals. How birds cope with the temporal geomagnetic variations caused by solar storms during migration is still unclear. We addressed this question by reproducing the effect of a solar storm on the geomagnetic field and monitoring the activity of three songbird species during autumn migration. We found that only the European robin reduced nocturnal migratory restlessness in response to simulated solar storms. At the same time, robins increased activity during early morning. We suggest that robins reduced activity at night when the perception of magnetic information would be strongly disrupted by temporal variations of the magnetic field, to extend their migration during daytime when several visual cues become available for orientation. The other two species, chiffchaff and dunnock, showing low or no nocturnal migratory activity, did not respond to the solar storm by changing activity.

Migratory state is not associated with differences in neural glucocorticoid or mineralocorticoid receptor expression in pine siskins

Although the endocrine system likely plays an important role in orchestrating the transition to a migratory state, the specific mechanisms by which this occurs remain poorly understood. Changes in glucocorticoid signaling are one proposed mechanism that may be important in migratory transitions. Although previous work has focused on the role of changes in circulating glucocorticoids, another potential mechanism is changes in the expression of its cognate receptors. Here, we test this hypothesis by comparing mRNA expression of the genes for the mineralocorticoid receptor (MR) and glucocorticoid receptor (GR) in two brain regions implicated in the regulation of migratory behavior (the hippocampus and hypothalamus) in pine siskins (Spinus pinus) sampled before or after the transition to a spring nomadic migratory state. Compared to pre-migratory birds, migratory birds had body conditions more indicative of physiological preparations for migration (e.g., larger body mass), and greater levels of nocturnal migratory restlessness. However, we found no differences between pre-migratory and migratory birds in the expression of GR or MR mRNA in either the hippocampus or hypothalamus. Thus, differences in expression of receptors for glucocorticoids do not appear to underly the observed differences in physiology and behavior across a migratory transition. Taken together with previous results showing no change in circulating corticosterone levels during this transition, our findings provide no evidence for a role of glucocorticoid signaling in the spring migratory transition of this species.