Compensatory behaviour after displacement in migratory birds

Using a meta-analysis approach we re-analysed orientation cage experiments with displaced migrants found in the literature. A rather large proportion of the orientation experiments showed directional shifts after displacements, indicating ability for birds to detect and react on such displacements. There was a clear difference between overcast and experiments where birds had a view of the starry sky. In experiments under a starry sky, the birds compensated the displacements, whilst under overcast unaltered or reverse orientations were generally displayed. This indicates a role for the stars to be involved in detection of the changes in position. Such a role of celestial cues is further stressed by the results of several studies manipulating a planetarium sky. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Plasticity versus repeatability in seabird migratory behaviour

Pelagic seabird populations can use several discrete wintering areas, but it is unknown if individuals use the same wintering area year after year. This would have consequences for their population genetic structure and conservation. We here study the faithfulness of individuals to a moulting area within and among years in a small pelagic seabird, the Thin-billed prion, which moult their primary feathers during the early part of the non-breeding period. According to stable carbon isotope ratios (δ¹³C) of these feathers, 90% of Thin-billed prions moult in Antarctic and 10% in South American waters. Repeated samples from individuals in 2 or 3 years indicated that several birds changed between Antarctic and South American moulting areas or vice versa. However, individuals moulting in an area in one year were more likely to do so again. Four out of five adults maintained highly conserved δ¹³C over the extended moulting period. One bird, however, had systematic changes in δ¹³C indicating latitudinal movements between the two areas during moult. Thus, the present data show that this seabird species has a highly flexible migratory strategy, not only at the population level, but also at the individual level, enabling these seabirds to exploit a highly unpredictable environment.     

Parasite transmission in a migratory multiple host system

A transmission model was devised for trichostrongyloid nematodes of saiga antelopes and domestic sheep in Kazakhstan. The framework extends previous models by including seasonal migration of saigas, contact with separate populations of sheep, and climate-driven stochasticity in herbage biomass and in the development, survival and migration onto herbage of free-living larvae. The model was parameterised for the contrasting life histories of Marshallagia, Haemonchus and Nematodirus, three important parasites of saigas and sheep in the region, and was successful at predicting broad qualitative patterns of infection dynamics in sheep and saigas. Parasite transmission between saigas and sheep was predicted to be most important for Marshallagia (from sheep to saigas in the south in winter, and onward transmission to sheep in the north in summer) and Haemonchus (from sheep in the north in summer via saigas to sheep further south in autumn). Model predictions for winter transmission of Marshallagia infection in saigas were consistent with field data, which showed that saigas culled before they have grazed the winter range carry lower burdens of this parasite than older animals. The model provides a mechanistic explanation for its predictions, which will assist hypothesis formation, and further the epidemiological basis of efforts to control parasite transmission between wildlife and livestock in both directions. A similar modelling approach could prove useful in other situations where detailed mechanistic models of parasite transmission are inappropriate in the face of parameter uncertainty and spatio-temporal variation in climate and host density. This is likely to include the majority of wildlife-parasite systems.     

Density-dependent migratory waves in the marine pelagic ecosystem

The migration of large aggregations of animals that sweep through the landscape is a phenomenon with large consequences in many ecosystems. It has been suggested that such migrations are mediated by resource depletion. Under this hypothesis it has been shown that simple foraging rules may generate density-dependent migratory waves (DDMW) in which the speed and amplitude increase with animal abundance. We tested these predictions on a 32-year data set of the spatial distribution of the two youngest age groups of a small pelagic schooling fish, the capelin (Mallotus villosus), by the end of their annual feeding migration in the Barents Sea. Our data suggest that the two age groups divided the Barents Sea by forming migratory waves that moved in opposite directions. The aggregation and spatial displacement of these waves increased with increasing age-specific abundance. However, possibly through social interactions, migratory pattern was modified by the abundance of the other age group.     

Wind selectivity of migratory flight departures in birds

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further. Received: 1 March 1999 / Received in revised form: 4 October 1999 / Accepted: 16 October 1999     

Delayed maturation of multiple signals in a migratory songbird

Many animals use multiple signals in sexual communication, but our understanding of the interactions between multiple signals, particularly in inexperienced breeders, is limited. In birds, delayed plumage maturation (DPM) is well documented; young birds appear duller than adults, despite reaching sexual maturity. Age-related changes in song structure are also common in songbirds, though the extent to which songs of yearling males differ from those of adults (delayed song maturation, DSM) and its prevalence in species with DPM is unknown. We tested for DSM in the mate-attraction (repeat) song of a species with dramatic DPM, the American redstart (Setophaga ruticilla). Repeat song structure of territorial yearling and adult males differed significantly, based on discriminant analysis of nine non-collinear song features. Combined with previous evidence of delayed maturation in territorial (serial) song, we provide the first evidence for DSM in different song types used in different behavioral contexts during the breeding season of a migratory songbird. Within adults, variation in repeat song was associated with pairing, earlier onset of breeding, and number of offspring sired, suggesting a potential benefit for expressing more adult-like song. We found no relationship between the expression of adult-like repeat song and plumage in either age class. These results indicate that delayed maturation of repeat song in yearling redstarts is not due to a generalized delay in signal development, and suggest further work is needed to determine whether mate attraction (for both yearling and adult males) provides greater pressure to sound more “adult-like” than to look more “adult-like”.     

Life history correlates of alternative migratory strategies in American Dippers

Partial migration is thought to be a critical step in the evolution of avian migration, but data on the life history correlates of alternative migratory strategies are extremely limited. We have studied a partially migratory population of American Dippers since 1999. This population is composed of sedentary individuals (residents) that maintain the same territory year round and altitudinal migrants that share winter grounds with residents, but move to higher elevations to breed. We used seven years of data on individually marked birds to (1) determine if individuals consistently use the same migratory strategy, (2) determine if offspring have the same strategy as their parents, and (3) estimate reproductive and survival rates of the two migratory strategies. We evaluate hypotheses for the persistence of partial migration and discuss their implication for the evolution of migration in sedentary populations. Individual American Dippers rarely switched migratory strategy (4/169 monitored more than one year). An individual's strategy, however, was not always that of its parents, indicating that, while migratory behavior may have a genetic component, environmental or social conditions probably influence the migratory strategy that an individual adopts. Sedentary dippers consistently had higher annual productivity (approximately 1.4 more fledglings/year) than migratory dippers, but mark-recapture models suggested that migratory dippers may have slightly higher survival than residents (approximately 3.4%). Migrants were estimated to have lower lifetime reproductive success than residents because their higher survival was insufficient to offset their lower productivity. Our data suggest that alternative migratory strategies in American Dippers are unlikely to be a fixed genetic dimorphism that persists because the two strategies have equal fitness, or because the relative fitness of the two strategies fluctuates over time. Migratory strategies in American Dippers are more likely to be condition dependent, and the two strategies persist because migrants "make the best of a bad job" by moving to higher elevations to breed. Because migrants obtained no fitness benefits by moving to seasonal breeding territories, our data are consistent with the hypothesis that migration could evolve in sedentary populations if competition for limited resources forces some individuals to seek breeding opportunities outside their initial range.     

Phenotype-dependent arrival time and its consequences in a migratory bird

Arrival times for migratory animals can be viewed as the result of an optimization process of costs and benefits of early arrival, and when the cost and benefit functions of early arrival depend on phenotypic quality, this will result in phenotype-dependent optimal arrival times. This hypothesis was tested for males of the migratory and sexually size-dimorphic barn swallow Hirundo rustica. The major cost of early arrival is poor environmental conditions which resulted in mortality of short-tailed early-arriving males in one year. The major benefits of early arrival are higher mating success, enhanced reproductive success, improved recruitment rates for offspring, and enhanced quality of the mate acquired. Annual variation in male arrival date is related to weather conditions at the breeding grounds, but also to some extent to weather conditions in the African winter quarters. Individual variation in arrival time can be explained by phenotype-dependent cost and benefit functions of early arrival. Male barn swallows with long tail ornaments arrived earlier than short-tailed males. The costs of early arrival should be particularly high under poor environmental conditions and lead to a stronger negative relationship between arrival date and phenotypic quality in years with poor environmental conditions. This prediction was confirmed by a stronger negative relationship between male tail length and date of arrival in years when arrival was relatively late because of poor weather. A female preference for early-arriving males may result in acquisition of good genes for optimal migratory behaviour, if migratory direction and extent have a genetic basis as shown in a number of different bird species.     

Group structure in a restricted entry system is mediated by both resident and joiner preferences

The benefits of grouping behaviour may not be equally distributed across all individuals within a group, leading to conflict over group membership among established group members, and between residents and outsiders attempting to join a group. Although the interaction between the preferences of joining individuals and existing group members may exert considerable pressure on group structure, empirical work on group living to date has focussed on free entry groups, in which all individuals are permitted entry. Using the humbug damselfish, Dascyllus aruanus, we examined a restricted entry grouping system, in which group residents control membership by aggressively rejecting potential new members. We found that the preferences shown by joining members were not always aligned with strategies that incurred the least harm from resident group members, suggesting a conflict between the preferences of residents and preferences of group joiners. Solitary fish preferred to join familiar groups and groups of size-matched residents. Residents were less aggressive towards familiar group joiners. However, resident aggression towards unfamiliar individuals depended on the size of the joining individual, the size of the resident and the composition of the group. These results demonstrate that animal group structure is mediated by both the preferences of joining individuals and the preferences of residents.     

Arrival timing and seasonal reproductive performance in a long-distance migratory landbird

The date when a landbird migrant arrives on its breeding grounds may have reproductive consequences. Generally, early arriving individuals begin breeding earlier and consequently experience greater seasonal reproductive performance. Here, we describe relationships between arrival timing and seasonal reproductive performance in the American redstart (Setophaga ruticilla), a long-distance passerine migrant, arriving at northerly breeding grounds in Michigans eastern Upper Peninsula. Evidence suggests that both males and females benefited from early arrival at the breeding grounds. Early males appeared to settle on higher quality territories and hatched nestlings sooner than later arrivals. Early females began their clutches early, produced heavier nestlings and possibly laid more eggs than later arrivals. Larger clutches and heavier offspring increase the likelihood of offspring recruiting into the breeding population. The findings of this study point to fitness consequences arising from when a bird arrives at its breeding grounds. These results also have implications for understanding how events occurring during spring migration influence reproductive performance as migratory delays likely influence arrival timing.Communicated by: W. Wiltschko     

Juvenile dispersal affects straying behaviors of adults in a migratory population

The resilience of organisms to large-scale environmental and climatic change depends, in part, upon the ability to colonize and occupy new habitats. While previous efforts to describe homing, or natal site fidelity, of migratory organisms have been hindered by the confounding effects of fragmented landscapes and management practices, realistic conservation efforts must include considerations of the behavioral diversity represented by animal movements and dispersal. Herein, we quantify straying away from natal origins by adult chinook salmon (Oncorhynchus tshawytscha) in a wild population that inhabits a pristine wilderness basin. Using natural isotopic signatures (7Sr/86Sr) to reconstruct the migratory behaviors of unhandled individuals over their entire life cycle, we identified ecological and behavioral factors influencing the propensity to stray. Our results indicate that natal site fidelity is scale dependent, ranging from 55% at -1-km distances to 87% at longer (> 10-km scale) distances, and juvenile dispersal and sex highly influence straying occurrence. These findings lend support for the conservation of behavioral diversity for population persistence, and we propose straying as a mechanism for maintaining genetic diversity at low population densities.     

Tidally-produced internal bands in the shell of Elminius modestus

Young specimens of Elminius modestus Darwin, were marked by treatment with calcium and strontium-enriched sea water, allowed to grow in the intertidal environment, and marked again. The number of growth bands laid down coincided with the number of tidal immersions. Strontium is not readily incorporated into the shell, even when its concentration in the external environment is greatly increased.     

Setting conservation priorities for migratory networks under uncertainty

Conserving migratory species requires protecting connected habitat along the pathways they travel. Despite recent improvements in tracking animal movements, migratory connectivity remains poorly resolved at a population level for the vast majority of species, thus conservation prioritization is hampered. To address this data limitation, we developed a novel approach to spatial prioritization based on a model of potential connectivity derived from empirical data on species abundance and distance traveled between sites during migration. We applied the approach to migratory shorebirds of the East Asian‐Australasian Flyway. Conservation strategies that prioritized sites based on connectivity and abundance metrics together maintained larger populations of birds than strategies that prioritized sites based only on abundance metrics. The conservation value of a site therefore depended on both its capacity to support migratory animals and its position within the migratory pathway; the loss of crucial sites led to partial or total population collapse. We suggest that conservation approaches that prioritize sites supporting large populations of migrants should, where possible, also include data on the spatial arrangement of sites.     

Asynchronous deposition of dense skeletal bands in Porites lutea

Alternate dense and less-dense skeletal bands in massive corals have been used for many years to record the history of growth in species such as Porites lutea and Montastrea annularis, based on the assumption that one dense band and one less-dense band is equivalent to a year's growth. This report demonstrates that specimens of Porites lutea Edwards and Haime (collected from the same neighbourhood in Phuket, Thailand, from November 1983 through November 1984) produce skeletal bands asynchronously and that one year's growth in corals from certain sites may regularly consist of four bands of varying density. The annual banding pattern observed at all sites includes the deposition of a dense band in response to high sedimentation loads and, probably, reduced light levels.     

Elevated testosterone advances onset of migratory restlessness in a nearctic-neotropical landbird

The primary cue for initiation of spring migratory restlessness (Zugunruhe) in landbird migrants is photoperiod. Gonadal hormones are known to have a role in the regulation of migratory disposition; however, the extent of their effect is not well understood. We examined the impact of exogenous testosterone on the onset of migratory restlessness in gray catbirds (Dumetella carolinensis). Catbirds were stratified by sex and randomly assigned to two rooms; individuals in one room were photoadvanced to initiate migratory restlessness and the other room was maintained on a 12:12 light/dark photoperiod. Each room had three groups (n?=?10/group); males with testosterone implants, males with empty implants, and females. We predicted that in the photoadvanced room males with testosterone implants would initiate migratory activity earlier than empty-implanted males. We found that in the photoadvanced group, testosterone-implanted males initiated migration 2 weeks prior to empty-implanted males, and 3 weeks prior to females. In the non-photoadvanced males, the testosterone-implanted males initiated migration at the same time as the corresponding group in the photoadvanced room, while the empty-implanted males and females did not exhibit Zugunruhe. Our results illustrate that elevated testosterone can advance the onset of Zugunruhe, even in the absence of an extended photoperiod. Additionally, the onset of migratory restlessness observed in the photoadvanced, non-testosterone-implanted males and females further supports the importance of photoperiod as a cue for migratory restlessness. An interesting observation was the intersexual differences in the onset of migratory activity in gray catbirds, a species not previously known to exhibit protandry.     

Geomagnetic field affects spring migratory direction in a long distance migrant

Night-migrating song birds travel to and from their wintering and breeding areas often separated thousands of kilometers apart and are clearly capable of finding intended goal areas from a distant location. Displacement experiments provide a useful way to highlight orientation and navigational skills in migrants. To investigate which cues birds actually use to compensate for displacement and the exact mechanism of each cue, experiments with manipulation of single cues are required. We conducted a simulated displacement of lesser whitethroats Sylvia curruca on spring migration. Birds were displaced not geographically but in geomagnetic space only, north and south of their breeding area to test whether they incorporate information from the geomagnetic field to find their breeding area. Lesser whitethroats held in southeast Sweden but experiencing a simulated displacement north of their breeding area (Norway) failed to show a consistent direction of orientation, whereas birds displaced south of their breeding area (Czech Republic) exhibited consistent northerly orientation, close to the expected seasonally appropriate direction, after displacement toward the trapping location. The absence of a clear compensatory direction in birds displaced north might be due to unfamiliar magnetic information or lack of sufficient information such as a magnetic gradient when moving around. By isolating one orientation cue, the geomagnetic field, we have been able to show that lesser whitethroats might incorporate geomagnetic field information to determine latitude during spring migration.     

Improved estimates of certainty in stable-isotope-based methods for tracking migratory animals.

The use of stable-hydrogen isotopes (deltaD) has become a common tool for estimating geographic patterns of movement in migratory animals. This method relies on broad and relatively predictable geographic patterning in deltaD values of precipitation, but these patterns are not estimated without error. In addition, deltaD measurements are relatively imprecise, particularly for organic tissue. Most models for estimating geographic locations have ignored these sources of error. Common modeling approaches include regression, range-matching, and likelihood-based assignment tests (including discriminant analysis). Here, we show the benefits of a simple stochastic extension to likelihood-based assignment tests that incorporates two estimable sources of error and describe the resulting influence on the certainty of assigning breeding origins for wintering American Redstarts (Setophaga ruticilla), a small Nearctic-Neotropical migratory bird. Through simulation, we incorporated both spatial interpolation error associated with models of deltaD in precipitation and analytical error associated with the measurement of deltaD in tissue samples. In general, assignments that did not include these sources of error fell within the ranges of the stochastic results, but the difference in proportion of birds assigned to any one breeding region varied by as much as 54%. To explore how the distribution of assignments generated from error models influenced the application of these results, we developed a simple model of winter habitat loss. We removed the proportion of Redstarts wintering at a particular site from the global population and then used the isotope-based assignments to predict the resulting population declines for each breeding region. This gave distributions of change in population sizes, some of which included no change or even a population increase. The sources of error we modeled may challenge the degree of certainty in the use of stable-isotope-based data on connectivity to predict population dynamics of migratory animals. We suggest that stronger inference will result from incorporating these sources of error into future studies that use deltaD or other stable isotopes to infer the geographic origin of individuals.     

Global population collapse in a superabundant migratory bird and illegal trapping in China

Persecution and overexploitation by humans are major causes of species extinctions. Rare species, often confined to small geographic ranges, are usually at highest risk, whereas extinctions of superabundant species with very large ranges are rare. The Yellow‐breasted Bunting (Emberiza aureola) used to be one of the most abundant songbirds of the Palearctic, with a very large breeding range stretching from Scandinavia to the Russian Far East. Anecdotal information about rapid population declines across the range caused concern about unsustainable trapping along the species’ migration routes. We conducted a literature review and used long‐term monitoring data from across the species’ range to model population trend and geographical patterns of extinction. The population declined by 84.3–94.7% between 1980 and 2013, and the species’ range contracted by 5000 km. Quantitative evidence from police raids suggested rampant illegal trapping of the species along its East Asian flyway in China. A population model simulating an initial harvest level of 2% of the population, and an annual increase of 0.2% during the monitoring period produced a population trajectory that matched the observed decline. We suggest that trapping strongly contributed to the decline because the consumption of Yellow‐breasted Bunting and other songbirds has increased as a result of economic growth and prosperity in East Asia. The magnitude and speed of the decline is unprecedented among birds with a comparable range size, with the exception of the Passenger Pigeon (Ectopistes migratorius), which went extinct in 1914 due to industrial‐scale hunting. Our results demonstrate the urgent need for an improved monitoring of common and widespread species’ populations, and consumption levels throughout East Asia.