Adverse wind conditions during northward Sahara crossings increase the in-flight mortality of Black-tailed Godwits

Long-distance migratory flights are predicted to be associated with higher mortality rates when individuals encounter adverse weather conditions. However, directly connecting environmental conditions experienced in-flight with the survival of migrants has proven difficult. We studied how the in-flight mortality of 53 satellite-tagged Black-tailed Godwits (Limosa limosa limosa) during 132 crossings of the Sahara Desert, a major geographical barrier along their migration route between The Netherlands and sub-Saharan Africa, is correlated with the experienced wind conditions and departure date during both southward and northward migration. We show that godwits experienced higher wind assistance during southward crossings, which seems to reflect local prevailing trade winds. Critically, we found that fatal northward crossings (15 deaths during 61 crossings) were associated with adverse wind conditions. Wind conditions during migration can thus directly influence vital rates. Changing wind conditions associated with global change may thus profoundly influence the costs of long-distance migration in the future

Individual Black-tailed Godwits do not stick to single routes:A hypothesis on how low population densities might decrease social conformity

The miniaturization of tracking devices is now rapidly increasing our knowledge on the spatiotemporal organization of seasonal migration. So far, most studies aimed at understanding within- and between-individual variation in migratory routines focus on single populations. This has also been the case for continental Black-tailed Godwits Limosa l. limosa (hereafter Godwits), with most work carried out on individuals from the Dutch breeding population, migrating in relatively large numbers in the westernmost part of the range. Here we report the migratory timing and routes of four adult individuals of the same subspecies from the low-density population in eastern Poland and compare this with previously published data on Godwits breeding in The Netherlands. During northward migration, the birds from Poland departed and arrived later from their wintering and breeding grounds. However, on southward migration the Polish breeding Godwits departed earlier, but arrived one month later than the Dutch birds on their wintering grounds in sub-Saharan Africa. Despite the small sample size of tracked birds from Poland, we find a significantly higher between-individual variation in timing during southward migration in Polish Godwits as compared to the Dutch Godwits. Furthermore, not only did migratory routes differ, but the few Polish Godwits tracked showed a higher level of between- and within-individual variation in route choice during both southward and northward migration. To explain this remarkable discrepancy, we propose that the properties of transmission of social information may be different between Godwits from a high-density population (i.e. the one in The Netherlands) and a low-density population (in Poland) and that this leads to different levels of canalization. To examine this hypothesis, future studies should not only follow individuals from an early age onwards, but also quantify and experimentally manipulate their social environments during migration

Migration route, stopping sites, and non-breeding destinations of adult Black-tailed Godwits breeding in southwest Fryslân, The Netherlands

In this paper, we extend our understanding of the migration of Black-tailed Godwits (Limosa limosa limosa) by describing: (1) the orientation and geographic locations of individual migratory routes and (2) the spatial distribution of godwits across seasons and years. We accomplish this using satellite-tracking data from 36 adult godwits breeding in the 200-ha Haanmeer polder in The Netherlands, from 2015 to 2018. During both southward and northward migration, godwits used a narrow migratory corridor along which most individuals made stops within a network of sites, especially the Bay of Biscay, France and Doñana, Spain. Most sites were used consistently by the same individuals across years. However, sites in Morocco were used during northward migration by 75% of individuals, but not revisited by the same individual across years. After southward migration, a small proportion (15%) of godwits spent the entire non-breeding period north of the Sahara, but most (85%) crossed the Sahara and spent at least part of the non-breeding season among seven coastal sites in West Africa and one site in the Inner Niger Delta. Although site-use patterns varied among individuals, individuals showed high site fidelity and were consistent in the number of sites they used from year to year. The considerable differences in the spatial distribution of individuals that breed within a kilometre of one another raise questions about the causes and consequences of individual migratory differences. We discuss that full annual cycle tracking of juveniles from birth to adulthood is needed to understand the source of these individual differences. Our results on the spatial distribution of godwits throughout their annual cycle lay an important foundation of information that can be used to help conserve this declining species

Variation in egg size of Black-tailed Godwits

As is the case for most avian species, there is considerable variation in the egg size of Continental Black-tailed Godwits Limosa l. limosa breeding in The Netherlands. It is interesting that egg size has costs and benefits yet varies considerably at the population level. To better understand this variation in egg size, we tested its relationship to a suite of individual and environmental factors. We found that egg size can decrease up to 2.8% throughout a breeding season and that egg size increases with clutch size by 1.4% with each additional egg in the clutch. Female body mass and body size explained 5% of the total variation in egg size observed across the population. Furthermore, females wintering south of the Sahara laid 3% smaller eggs than those wintering north of the Sahara. We also found that egg size increases with age, which may indicate age-related differences in the endogenous and/or exogenous conditions of females. The variation in egg size was, however, mostly the result of consistent differences among individuals across years (repeatability = 0.60). A comparison of daughters with mothers suggested that most of this individual repeatability reflects heritable variation (heritability = 0.64). The actual individual traits that underlie this heritable variation among individuals remain mostly undetermined. Smaller eggs did have a slightly lower chance of hatching, but we found no relationship between egg size and chick survival. Finally, nest and chick survival were strongly correlated with lay date. Thus, in Black-tailed Godwits, lay date may actually reflect a female's endogenous and/or exogenous condition at the moment of egg-laying. This finding may be general across birds, since food supplementation experiments usually result in advanced laying and larger clutch sizes rather than in larger eggs

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

High Migratory Survival and Highly Variable Migratory Behavior in Black-Tailed Godwits

Few studies have been able to directly measure the seasonal survival rates of migratory species or determine how variable the timing of migration is within individuals and across populations over multiple years. As such, it remains unclear how likely migration is to affect the population dynamics of migratory species and how capable migrants may be of responding to changing environmental conditions within their lifetimes. To address these questions, we used three types of tracking devices to track individual black-tailed godwits from the nominate subspecies (Limosa limosa limosa) throughout their annual cycles for up to 5 consecutive years. We found that godwits exhibit considerable inter- and intra-individual variation in their migratory behavior across years. We also found that godwits had generally high survival rates during migration, although survival was reduced during northward flights across the Sahara Desert. These patterns differ from those observed in most other migratory species, suggesting that migration may only be truly dangerous when crossing geographic barriers that lack emergency stopover sites and that the levels of phenotypic flexibility exhibited by some populations may enable them to rapidly respond to changing environmental conditions

Warming springs and habitat alteration interact to impact timing of breeding and population dynamics in a migratory bird

In seasonal environments, increasing spring temperatures lead many taxa to advance the timing of reproduction. Species that do not may suffer lower fitness. We investigated why black-tailed godwits (Limosa limosa limosa), a ground-breeding agricultural grassland shorebird, have not advanced timing of reproduction during the last three decades in the face of climate change and human-induced habitat degradation. We used data from an 11-year field study to parameterize an Integral Projection Model to predict how spring temperature and habitat quality simultaneously influence the timing of reproduction and population dynamics. We found apparent selection for earlier laying, but not a correlation between the laying dates of parents and their offspring. Nevertheless, in warmer springs, laying dates of adults show a stronger positive correlation with laying date in previous springs than in cooler ones, and this leads us to predict a slight advance in the timing of reproduction if spring temperatures continue to increase. We also show that only in landscapes with low agricultural activity, the population can continue to act as a source. This study shows how climate change and declining habitat quality may enhance extinction risk

Individual Black-Tailed Godwits Do Not Stick to Single Routes: A Hypothesis on How Low Population Densities Might Decrease Social Conformity

The miniaturization of tracking devices is now rapidly increasing our knowledge on the spatiotemporal organization of seasonal migration. So far, most studies aimed at understanding within- and between-individual variation in migratory routines focus on single populations. This has also been the case for continental Black-tailed Godwits Limosa l. limosa (hereafter Godwits), with most work carried out on individuals from the Dutch breeding population, migrating in relatively large numbers in the westernmost part of the range. Here we report the migratory timing and routes of four adult individuals of the same subspecies from the low-density population in eastern Poland and compare this with previously published data on Godwits breeding in The Netherlands. During northward migration, the birds from Poland departed and arrived later from their wintering and breeding grounds. However, on southward migration the Polish breeding Godwits departed earlier, but arrived one month later than the Dutch birds on their wintering grounds in sub-Saharan Africa. Despite the small sample size of tracked birds from Poland, we find a significantly higher between-individual variation in timing during southward migration in Polish Godwits as compared to the Dutch Godwits. Furthermore, not only did migratory routes differ, but the few Polish Godwits tracked showed a higher level of between- and within-individual variation in route choice during both southward and northward migration. To explain this remarkable discrepancy, we propose that the properties of transmission of social information may be different between Godwits from a high-density population (i.e. the one in The Netherlands) and a low-density population (in Poland) and that this leads to different levels of canalization. To examine this hypothesis, future studies should not only follow individuals from an early age onwards, but also quantify and experimentally manipulate their social environments during migration

Data_Capture_Recapture_Survival_Analysis

Capture-recapture dataset used for the survival analysis. CR-history contains two digits per year, first digit is only used when individual was ringed as chick. Further details explained in file