Light-level Geolocators Research Articles

Drivers of variation in migration behavior for a linked population of long-distance migratory passerine

AbstractDespite advances in tracking technologies, migration strategies remain poorly studied for many small-bodied passerines. Understanding variation within a migration strategy is important as variation impacts a population’s resilience to environmental change. Timing, pathway, and stopovers vary based on intrinsic and extrinsic factors that impact individual migration decisions and capacity. Here, we studied drivers of variation in migration across a linked population of Golden-winged Warbler (Vermivora chrysoptera) using data from 37 light-level geolocators. We tested if behaviors vary in response to extrinsic factors: season, year, and proximity to a large geographic barrier—the Gulf of Mexico—and intrinsic factors: age and wing chord. Spring migration was nearly twice as fast as fall migration, with tightly correlated arrival and departure dates that were consistent among years, in contrast to no correlation or consistency in fall. This aligns with predictions for selection to minimize time spent migrating in spring and a relaxation of that pressure in fall. Twenty-nine birds staged for multiple days (mean: 7.5, SE: 0.6) in stopover habitats before crossing the Gulf of Mexico in spring, but 6 individuals overwintering closer to the Gulf coast forewent the stopover and completed migration 8 days faster. These findings suggest birds capable of crossing the Gulf without a stopover may experience a selective advantage by minimizing total migration time. After crossing the Gulf, individuals reduced travel speed and stopover duration, indicating constraints on movement differ before and after the barrier. Wing chord, but not age, positively predicted the total distance and duration of migration, and neither varied with timing, suggesting migration distance impacts morphology, but strategies do not vary with age. Ultimately, we find undescribed stopover locations south of the Gulf are important for most of the population, while high variation in migration behaviors suggest potential resilience to changing environmental conditions.Causantes de variación en el comportamiento migratorio para una población vinculada de un paseriforme migratorio de larga distancia

Sex-Specific Spatiotemporal Variation and Carry-Over Effects in a Migratory Alpine Songbird

For migratory animals, events at one stage of the annual cycle can produce constraints or benefits that carry over to subsequent stages. Differing life-history strategies among individuals can influence the expression of these carry-over effects, leading to pronounced within-population variation in migration. For example, reproductive roles can drive spatiotemporal segregation during the non-breeding season and promote sex-specific carry-over effects, such as reproductive effort affecting autumn migration behaviour. For an alpine breeding population of horned larks Eremophila alpestris in northern British Columbia, Canada, we addressed sex-specific variation in migration behaviour and carry-over effects during both autumn and spring migration using light-level geolocators. Males spent more time farther north and arrived an average of 6 days earlier at the breeding site in spring. Females delayed autumn departure following greater reproductive effort, in turn demonstrating flexible migration behaviour by increasing migration speed and decreasing stopover use. Males maintained autumn migration behaviour regardless of reproductive effort or departure date. Finally, both sexes used staging areas in spring (average stopover = 41 days), with consequences for breeding success. Individuals that used staging areas during spring migration exhibited greater nest success and produced 1.8 more fledglings on average than those that migrated directly from their winter site. Consistent use of staging areas may allow individuals to monitor environmental conditions and optimize their breeding arrival date to acquire high-quality territories while avoiding the cost of arriving too early in a harsh alpine habitat. Overall, our results indicate: (1) sex-specific flexibility in migration strategies that carry-over to and from the reproductive period, and (2) spring staging areas may be a critical component of the annual life-cycle for alpine breeding larks. These behaviours may be particularly important for alpine and arctic birds because the stochastic nature of their breeding habitat likely selects for flexible responses to prevailing conditions.

No detectable effect of light-level geolocators on the behaviour and fitness of a long-distance migratory seabird

Light-level geolocators are increasingly popular devices for tracking migratory birds. However, to ensure that data on migratory behaviour represent natural behaviour that is not affected by potentially harmful effects of carrying such a device, their effects on behaviour and fitness should be assessed. A review of studies that tested for effects of tarsus-mounted light-level geolocators on seabirds showed that results are equivocal and often difficult to interpret due to the inclusion of only few traits and/or the lack of a proper experimental design. We therefore experimentally tested whether tarsus-mounted light-level geolocators affected a long-distance migratory seabird, the Common Tern Sterna hirundo. Using a well-matched treatment and control group, including both males and females, we tested whether light-level geolocators, deployed in the second half of incubation, affected the subsequent share of incubation, provisioning rate, reproductive performance, phenology or survival of tagged birds or their partners. In the year of deployment, we found no evidence for the behaviour of tagged birds or their partners to be affected by the geolocators. Moreover, we found no effect on their reproductive performance and departure date from the breeding colony. Finally, neither local survival to the next season, nor arrival date to the breeding colony in that season differed between tagged birds or their partners and control birds. These results suggest that a year of carrying a light-level geolocator attached to the tarsus has negligible effects on Common Terns and that such a device can be used to study their migratory behaviour without causing problems or introducing bias.

Migration pattern of Gambel’s White-crowned Sparrow along the Pacific Flyway

White-crowned Sparrow (Zonotrichia leucophrys) populations of western North America exhibit dramatic differences in life history strategies including migration behavior. However, individual migration strategies and population-level migratory patterns remain largely unknown for this species. Here, we focused on the long-distance migratory subspecies, Gambel’s White-crowned Sparrow (Zonotrichia leucophrys gambelii). We used ringing, tracking and stable hydrogen isotope (δ2H) analysis of individuals migrating along the Pacific Flyway to assess individual phenology and routes as well as the pattern of connectivity between breeding and non-breeding sites. Results from all three methods, consisting of 79 ring recoveries, four light level geolocator tracks and 388 feather δ2H values, indicate low degrees of migratory connectivity. The isotope data provide evidence for leapfrog migration with the more southerly populations traveling greater distances to the breeding grounds than more centrally wintering individuals. Location estimates of four annual journeys revealed individually consistent migration strategies with relatively short flight bouts separated by two to three and two to six stopover sites during spring and autumn migration, respectively. However, combined results from all methods indicate high variability in migration distance among individuals. These findings confirm the phenotypic flexibility observed within this species and highlight the potential of White-crowned Sparrows for further investigations of evolutionary adaptations to ongoing changes in the environment.

An ontogenetic perspective on migratory strategy of a long‐lived pelagic seabird: Timings and destinations change progressively during maturation

The processes that drive the ontogeny of migratory strategies in long-lived animals with slow maturation remain enigmatic. While some short-lived migrants are known or believed to repeat the same migratory patterns throughout their lives, little is known on the time required for immature long-lived migrants to progressively acquire adult-like migratory behaviours, or which aspects take longer to refine during the maturation process. Here, we studied the ontogeny of long-distance migratory strategies and related patterns of spatial distribution in a long-lived seabird species during the annual cycle. To do so, we deployed light-level geolocators on 4- to 9-year-old immature Cory's shearwaters (Calonectris borealis) and on breeding adults. We revealed that migratory timings and destinations of young shearwaters progressively changed with age. The effect of ageing was remarkably evident on spring migratory performance and phenology. Birds gradually shortened the duration of the non-breeding period by advancing departure date and reducing travelling time, which resulted in a sequential arrival at the colony of the various age contingents. Ageing immatures gradually changed from a more exploratory strategy to a more conservative way of exploiting resources, reducing both their year-round spatial spread across oceanic domains and the total distance travelled. Immatures always performed a trans-equatorial migration to the Southern Hemisphere, contrasting with 17% of the adults which remained in the North Atlantic year-round. Finally, during the breeding season immatures were widely dispersed through the North Atlantic reducing their overlap with breeding adults. Our long-term study provides empirical support to the hypothesis that in long-lived species, the refinement of migratory behaviour and year-round spatial distribution is a progressive process mediated by age and experience, where life stage constraints and competition for resources may also play a role. The emerging pattern suggests that for some avian taxa, the ontogeny of migratory strategy is a prolonged, complex and dynamic process.

Light-level geolocator analyses: A user's guide.

Light-level geolocator tags use ambient light recordings to estimate the whereabouts of an individual over the time it carried the device. Over the past decade, these tags have emerged as an important tool and have been used extensively for tracking animal migrations, most commonly small birds. Analysing geolocator data can be daunting to new and experienced scientists alike. Over the past decades, several methods with fundamental differences in the analytical approach have been developed to cope with the various caveats and the often complicated data. Here, we explain the concepts behind the analyses of geolocator data and provide a practical guide for the common steps encompassing most analyses - annotation of twilights, calibration, estimating and refining locations, and extraction of movement patterns - describing good practices and common pitfalls for each step. We discuss criteria for deciding whether or not geolocators can answer proposed research questions, provide guidance in choosing an appropriate analysis method and introduce key features of the newest open-source analysis tools. We provide advice for how to interpret and report results, highlighting parameters that should be reported in publications and included in data archiving. Finally, we introduce a comprehensive supplementary online manual that applies the concepts to several datasets, demonstrates the use of open-source analysis tools with step-by-step instructions and code and details our recommendations for interpreting, reporting and archiving.

Reconstruction of long-distance bird migration routes using advanced machine learning techniques on geolocator data.

Geolocators are a well-established technology to reconstruct migration routes of animals that are too small to carry satellite tags (e.g. passerine birds). These devices record environmental light-level data that enable the reconstruction of daily positions from the time of twilight. However, all current methods for analysing geolocator data require manual pre-processing of raw records to eliminate twilight events showing unnatural variation in light levels, a step that is time-consuming and must be accomplished by a trained expert. Here, we propose and implement advanced machine learning techniques to automate this procedure and we apply them to 108 migration tracks of barn swallows ( Hirundo rustica). We show that routes reconstructed from the automated pre-processing are comparable to those obtained from manual selection accomplished by a human expert. This raises the possibility of fully automating light-level geolocator data analysis and possibly analysing the large amount of data already collected on several species.

Nonbreeding season movements of a migratory songbird are related to declines in resource availability

AbstractThere have been an increasing number of observations of itinerancy in migratory songbirds, where individuals move among 2 or more widely separated areas during the “stationary” nonbreeding season. Knowledge of such movements and an understanding of what drives them are important for predicting how migratory populations will respond to environmental change. In this study, we investigated nonbreeding movements of the Tree Swallow (Tachycineta bicolor), an aerial insectivore that breeds across North America and spends the nonbreeding season around the Gulf of Mexico, Florida, Mexico, Central America, and the Caribbean. With year-round tracking data obtained from 133 light-level geolocators deployed at 12 breeding sites ranging from Alaska to Nova Scotia to North Carolina, we show that 44% of individuals made at least one large-scale movement (range: 301–1,744 km) within the nonbreeding range. The frequency of itinerancy decreased with longitude, such that 75% of individuals made a movement in the western portion of the nonbreeding range compared to only 31% in the east. Using the Normalized Difference Vegetation Index (NDVI) as a proxy for resource availability, we found that when individuals did move, they were more likely to move from sites where resources were deteriorating faster (a more negative change in NDVI prior to departure) than their destination sites. There was also evidence that individuals moved to destination sites with higher NDVI and temperature in the autumn, but not in the winter. Our results suggest movements of Tree Swallows during the nonbreeding season are influenced by resource availability, but because not all individuals used multiple nonbreeding sites, the density of individuals at a site and the level of competition may have also been a factor influencing nonbreeding season movements.

Concentration of a widespread breeding population in a few critically important nonbreeding areas: Migratory connectivity in the Prothonotary Warbler

AbstractOne of the greatest challenges to informed conservation of migratory animals is elucidating spatiotemporal variation in distributions. Without such information, it is impossible to understand full-annual-cycle ecology and effectively implement conservation actions that address where and when populations are most limited. We deployed and recovered light-level geolocators (n = 34) at 6 breeding sites in North America across the breeding range of a declining long-distance migratory bird, the Prothonotary Warbler (Protonotaria citrea). We sought to determine migratory routes, stopover location and duration, and the location of overwintering grounds. We found that the species exhibits a large-scale, east‒west split in migratory routes and weak migratory connectivity across its range. Specifically, almost all individuals, regardless of breeding origin, overlapped in their estimated wintering location in northern Colombia, in an area 20% the size of the breeding range. Additionally, most of the individuals across all breeding locations concentrated in well-defined stopover locations in Central America while en route to Colombia. Although error inherent in light-level geolocation cannot be fully ruled out, surprisingly much of the estimated wintering area included inland areas even though the Prothonotary Warbler is considered a specialist on coastal mangroves in winter. Based on these results, conservation efforts directed at very specific nonbreeding geographical areas will potentially have benefits across most of the breeding population. Our findings highlight the importance of using modern technologies to validate assumptions about little-studied portions of a species’ annual cycle, and the need to distribute sampling across its range.

A Migratory Divide Among Red-Necked Phalaropes in the Western Palearctic Reveals Contrasting Migration and Wintering Movement Strategies

Non-breeding movement strategies of migratory birds may be expected to be flexibly adjusted to the distribution and quality of habitat, but only few studies compare movement strategies between populations using distinct migration routes and wintering areas. In thisour study, individual movement strategies of Rred-necked pPhalaropes Phalaropus lobatus, a long-distance migratory wader using saline waters in the non-breeding period, were studied using light-level geolocators. Results revealed the existence of two populations with distinct migration routes and wintering areas: one breeding in the north-eastern North Atlantic and migrating ca. 10,000 km oversea to the tropical eastern Pacific Ocean and the other breeding in Fennoscandia and Russia migrating ca. 6,000 km – largely over land – to the Arabian Sea (Indian Ocean). In line with our expectations, the transoceanic migration between the North Atlantic and the Pacific was associated with proportionately longer wings, a more even spread of stopovers in autumn and a higher migration speed in spring compared to the migration between Fennoscandian-Russian breeding grounds and the Arabian Sea. In the wintering period, birds wintering in the Pacific were stationaryresided in roughly a singlethe same area, whereas individuals wintering in the Arabian Sea showed individually consistent movementsd extensively between different areas, reflecting differences in spatio-temporal variation in primary productivity between the two wintering areas. Our study is unique in showing how habitat distribution shapes movement strategies over the entire non-breeding period within a species.

Weak effects of geolocators on small birds: A meta-analysis controlled for phylogeny and publication bias.

Currently, the deployment of tracking devices is one of the most frequently used approaches to study movement ecology of birds. Recent miniaturization of light-level geolocators enabled studying small bird species whose migratory patterns were widely unknown. However, geolocators may reduce vital rates in tagged birds and may bias obtained movement data. There is a need for a thorough assessment of the potential tag effects on small birds, as previous meta-analyses did not evaluate unpublished data and impact of multiple life-history traits, focused mainly on large species and the number of published studies tagging small birds has increased substantially. We quantitatively reviewed 549 records extracted from 74 published and 48 unpublished studies on over 7,800 tagged and 17,800 control individuals to examine the effects of geolocator tagging on small bird species (body mass <100g). We calculated the effect of tagging on apparent survival, condition, phenology and breeding performance and identified the most important predictors of the magnitude of effect sizes. Even though the effects were not statistically significant in phylogenetically controlled models, we found a weak negative impact of geolocators on apparent survival. The negative effect on apparent survival was stronger with increasing relative load of the device and with geolocators attached using elastic harnesses. Moreover, tagging effects were stronger in smaller species. In conclusion, we found a weak effect on apparent survival of tagged birds and managed to pinpoint key aspects and drivers of tagging effects. We provide recommendations for establishing matched control group for proper effect size assessment in future studies and outline various aspects of tagging that need further investigation. Finally, our results encourage further use of geolocators on small bird species but the ethical aspects and scientific benefits should always be considered.

Variation From an Unknown Source: Large Inter-individual Differences in Migrating Black-Tailed Godwits

Variation in migratory behavior is the result of different individual strategies and fluctuations in individual performances. A first step toward understanding these differences in migratory behavior among individuals is, therefore, to assess the relative contributions of inter- and intra-individual differences to this variation. We did this using light-level geolocators deployed on the breeding grounds to follow continental black-tailed godwits (Limosa limosa limosa) throughout their south- and northward migrations over multiple years. Based on repeated tracks from 36 individuals, we found two general patterns in godwit migratory behavior: First, migratory timing in black-tailed godwits varies mostly because individual godwits migrate at different times of the year. Second, individuals also exhibit considerable variation in timing within their respective migratory windows. Although the absolute amount of inter-individual variation in timing decreased over the course of northward migration, individual godwits still arrived at their breeding grounds across a span of more than 5 weeks. These differences in migratory timing among individuals are larger than those currently observed in other migratory bird species and suggest that the selective forces that limit the variation in migratory timing in other species are relaxed or absent in godwits. Furthermore, we could not attribute these individual differences to the sex or wintering location of an individual. We suggest that different developmental trajectories enabled by developmental plasticity likely result in these generally consistent, life-long annual routines. To investigate this possibility and to gain an understanding of the different selection pressures that could be acting during migration and throughout a godwit's life, future studies should track juvenile godwits and other migratory birds from birth to adulthood while also manipulating their spatiotemporal environment during development.

A range-wide domino effect and resetting of the annual cycle in a migratory songbird.

Latitudinal differences in timing of breeding are well documented but how such differences carry over to influence timing of events in the annual cycle of migratory birds is not well understood. We examined geographical variation in timing of events throughout the year using light-level geolocator tracking data from 133 migratory tree swallows ( Tachycineta bicolor) originating from 12 North American breeding populations. A swallow's breeding latitude influenced timing of breeding, which then carried over to affect breeding ground departure. This resulted in subsequent effects on the arrival and departure schedules at autumn stopover locations and timing of arrival at non-breeding locations. This 'domino effect' between timing events was no longer apparent by the time individuals departed for spring migration. Our range-wide analysis demonstrates the lasting impact breeding latitude can have on migration schedules but also highlights how such timing relationships can reset when individuals reside at non-breeding sites for extended periods of time.

Geolocator deployment reduces return rate, alters selection, and impacts demography in a small songbird.

In the past few years, miniature light-level geolocators have been developed for tracking wild bird species that were previously too small to track during their full annual cycle. Geolocators offer an exciting opportunity to study the full annual cycle for many species. However, the potential detrimental effects of carrying geolocators are still poorly understood, especially for small-bodied birds. Here, we deployed light-level geolocators on common yellowthroat warblers (Geothlypis trichas). Over two years, we monitored return rates and neighborhood demography for 40 warblers carrying a geolocator and 20 reference birds that did not carry a geolocator. We compared the two groups with long-term data from 108 unmanipulated birds breeding at the same location in previous and subsequent years. Overall, we found that individuals carrying a geolocator were less likely to return to the study site in the following year (21% to 33% returned, depending on inclusion criteria) than either contemporaneous controls (55%) or long-term controls (55%). Among birds marked with geolocators, we also detected viability selection for greater wing length, whereas this pattern was not present in control birds. Finally, in each year after geolocator deployment, inexperienced breeders colonized vacant territories and this demographic effect persisted for two years after deployment. Sexual selection and ornamentation are strongly age-dependent in this system, and behavioral data collected after geolocator deployment is likely to differ systematically from natural conditions. Clearly geolocators will continue to be useful tools, but we suggest that future studies should carefully consider the potential for biased returns and the ecological validity of behavioral data collected from geolocator marked populations.

Identifying migration routes and non-breeding staging sites of adult males of the globally threatened Aquatic WarblerAcrocephalus paludicola

SummaryBirds that are long-distance migrants partition their annual cycle among a number of locations over a large spatial range. The conservation of these species is particularly complex because it requires attention to a number of different and distant habitats based on knowledge of migratory phenology, routes and staging areas. In the case of the globally threatened Aquatic WarblerAcrocephalus paludicola, a habitat specialist that breeds in Europe and spends the boreal winter in sub-Saharan Africa, non-breeding staging areas were widely unknown until recently. We applied light-level geolocators to adult male Aquatic Warblers at breeding sites in Belarus and Ukraine. Data from eight retrieved geolocators confirmed a south-west and then westward migration route through Europe via the northern Mediterranean with staging sites on the Iberian Peninsula and occasionally France and north-west Africa. In sub-Saharan Africa, final staging areas were in Mali, either in the Inner Niger Delta or at (presumably) small water bodies in the desert, with one bird most likely staying in the north of Ivory Coast south of the previously assumed range. The birds probably stayed in the final staging areas for most of the non-breeding season, but the logger on only one bird provided data throughout the entire migration cycle. Pre-nuptial migration of the latter was by a more easterly route than the southward migration. Our study suggests that conservation strategies for Aquatic Warbler north of the Sahara should include consideration of unknown staging sites in the northern Mediterranean as well as on the Iberian Peninsula and in north-west Africa. South of the Sahara, our study demonstrates the importance of the Sahel for the conservation of the Aquatic Warbler, including both the major floodplains of the Niger River and small Sahelian wetlands that are under pressure from human development.

Annual GPS tracking reveals unexpected wintering area in a long-distance migratory songbird

Recent technological development has made it possible to pinpoint precise locations of small migratory songbirds throughout their annual cycle, providing the opportunity for improving our understanding of year-round habitat use. Here, we use GPS loggers to map the exact location and habitat use at stationary sites throughout the annual cycle of a long-distance migratory songbird, the Red-backed Shrike Lanius collurio. Although the main staging sites confirmed previous findings from light-level geolocation studies, one individual wintered in south-western Chad, an area with only a few historical records of this species. This study highlights opportunities for answering new questions and gaining more knowledge using fine-scale tracking of migratory songbirds.

A quantitative ring recovery analysis of migration in European Water Rails Rallus aquaticus

The Water Rail Rallus aquaticus is generally regarded as a facultative migrant in which migration patterns vary geographically. However, quantitative analyses of migration directions and how migration distances vary across Europe are hitherto lacking. We analysed recoveries of Water Rails ringed in Europe and tested for a general migration direction in this material. From previous literature descriptions we predicted birds to migrate farther the longer north and east they breed. More than half of the birds included in the data set were ringed in Germany (48%) and Hungary (12%) and most winter recoveries came from SW coastal Europe. We found that the migratory direction in autumn was strongly oriented towards SW with no effect of latitude or longitude of the breeding site. There were no differences in migration direction between old (2Y+) and young (1Y) birds, or between males and females, although sample sizes were admittedly small in the latter comparison. As predicted, migration distances were positively correlated with both latitude and longitude of breeding sites. We encourage more trapping and ringing of Water Rails breeding and wintering across Europe in order to improve the ring recovery data. The use of modern tracking devices like light-level geolocators also has a high potential of improving our understanding of migratory ecology in this secretive species.

Ten years tracking the migrations of small landbirds: Lessons learned in the golden age of bio-logging

ABSTRACT In 2007, the first miniature light-level geolocators were deployed on small landbirds, revolutionizing the study of migration. In this paper, we review studies that have used geolocators to track small landbirds with the goal of summarizing research themes and identifying remaining important gaps in understanding. We also highlight research and opportunities using 2 recently developed tracking technologies: archival GPS tags and automated radio-telemetry systems. In our review, we found that most (54%) geolocator studies focused on quantifying natural history of migration, such as identifying migration routes, nonbreeding range, and migration timing. Studies of behavioral ecology (20%) uncovered proximate drivers of movements, including en route habitat quality; that migration routes, but not timing, may be flexible in some species; and different age and sex classes show significant differences in migration strategy. Studies of the evolution of migration (9%) have illustrated that migration is a ...

Light-level geolocation in polar regions with 24-hour daylight

Light-level geolocation in polar regions with 24-hour daylight

Follow the rain? Environmental drivers ofTyrannusmigration across the New World

ABSTRACT Predictable seasonal changes in resources are thought to drive the timing of annual animal migrations; however, we currently understand little about which environmental cues or resources are tracked by different migratory bird species across the planet. Understanding which environmental cues or resources birds track in multiple migratory systems is a prerequisite to developing generalizable conservation plans for migratory birds in a changing global environment. Within the New World, climatic differences experienced by Nearctic–Neotropical migratory (NNM; i.e. breed in North America and spend the nonbreeding period in the Neotropics) and Neotropical austral migratory (NAM; i.e. breed and spend the nonbreeding period wholly within South America) bird species suggest that their migratory strategies may be shaped by unique selective pressures. We used data gathered from individuals fitted with light-level geolocators to build species distribution models (SDMs) to test which environmental factors dri...