Spatial and temporal variation in population trends in a long‐distance migratory bird

Aim Over the past three decades, evidence has been growing that many Afro‐Palaearctic migratory bird populations have suffered sustained and severe declines. As causes of these declines exist across both the breeding and non‐breeding season, identifying potential drivers of population change is complex. In order to explore the roles of changes in regional and local environmental conditions on population change, we examine spatial and temporal variation in population trajectories of one of Europe’s most abundant Afro‐Palaearctic summer migrants, the willow warbler, Phylloscopus trochilus. Location Britain and Ireland. Methods We use national survey data from Britain and Ireland (BBS: BTO/RSPB/JNCC Breeding Bird Survey and CBS: BWI/NPWS/Heritage Council Countryside Breeding Survey) from 1994 to 2006 to model the spatial and temporal variation in willow warbler population trends. Results Across Britain and Ireland, population trends follow a gradient from sharp declines in the south and east of England to shallow declines and/or slight increases in parts of north and west England, across Scotland and Ireland. Decreasing the spatial scale of analysis reveals variation in both the rate and spatial extent of population change within central England and the majority of Scotland. The rates of population change also vary temporally; declines in the south of England are shallower now than at the start of the time series, whereas populations further north in Britain have undergone periods of increase and decline. Main conclusion These patterns suggest that regional‐scale drivers, such as changing climatic conditions, and local‐scale processes, such as habitat change, are interacting to produce spatially variable population trends. We discuss the potential mechanisms underlying these interactions and the challenges in addressing such changes at scales relevant to migratory species.     

Annual variation in long‐distance dispersal driven by breeding and non‐breeding season climatic conditions in a migratory bird

Long‐distance dispersal is a fundamental process in ecology and evolution but the factors that influence these movements remain poorly understood in most species. We used stable hydrogen isotopes to quantify the rate and direction of long‐distance immigration in a breeding population of American redstarts and to test whether the settlement decisions that result in long‐distance dispersal are driven by habitat saturation or by the phenology of breeding‐season resources. Our results provide evidence that both natal dispersal and breeding dispersal were influenced by the timing of breeding‐season phenology, with both age classes more likely to disperse north in years when the onset of breeding‐season phenology occurs earlier than normal. Yearlings were also more likely to disperse north following winters with poor habitat quality on their non‐breeding grounds, demonstrating that carry‐over effects from the non‐breeding season influence natal dispersal in this species. Collectively, these results are consistent with the hypothesis that American redstarts use the phenology of breeding season resources as a cue to select breeding sites. Our results suggest that long‐distance dispersal may allow individuals to rapidly respond to advancing phenology caused by global climate change, though their ability to do so may be constrained by long‐term decline in habitat quality predicted for their tropical non‐breeding grounds.     

Form,function and consequences of density dependence in a long‐distance migratory bird

The density dependence of demographic parameters and its implications for population regulation have long been recognized. Recent work has revealed potential effects of density on mating systems and sexual selection, but few studies concurrently assess the consequences of density on both demography and sexual selection. Such an approach is important because population processes and individual behaviors can interact to influence population growth and evolutionary trajectories. In this study, we tested the density dependence of breeding success, extra‐pair paternity, and the opportunity for sexual selection in a population of American redstarts Setophaga ruticilla using two different measures of density. To evaluate temporal patterns, we analyzed annual territory density, based on the total number of territories at our study site each year. To evaluate spatial patterns, we analyzed local territory density within years, based on the number of territories surrounding a focal territory. Greater annual density was associated with fewer offspring fledged per female, a reduced mean population rate of fledging success, and a lower relative contribution of extra‐pair paternity to male fitness. Greater local density was associated with fewer offspring fledged, reduced fledgling success, higher rates of nest loss, and higher rates of paternity loss on focal territories. Interestingly, greater local density was also associated with greater nestling mass on focal territories, which could imply that more densely‐packed territories contain superior resources. Overall, our results suggest that the effects of crowding via greater territory density reduce fecundity through increased nest predation, rather than reduced food availability, and increase rates of extra‐pair paternity. Thus, the selective pressures faced by individuals and their reproductive behaviors are likely to differ based on the annual and local density they experience, which may then feed back into population demography.     

The large‐scale drivers of population declines in a long‐distance migratory shorebird

Migratory species can travel tens of thousands of kilometers each year, spending different parts of their annual cycle in geographically distinct locations. Understanding the drivers of population change is vital for conserving migratory species, yet the challenge of collecting data over entire geographic ranges has hindered attempts to identify the processes leading to observed population changes. Here, we use remotely sensed environmental data and bird count data to investigate the factors driving variability in abundance in two subspecies of a long‐distance migratory shorebird, the bar‐tailed godwit Limosa lapponica. We compiled a spatially and temporally explicit dataset of three environmental variables to identify the conditions experienced by each subspecies in each stage of their annual cycle (breeding, non‐breeding and staging). We used a Bayesian N‐mixture model to analyze 18 years of monthly count data from 21 sites across Australia and New Zealand in relation to the remote sensing data. We found that the abundance of one subspecies L. l. menzbieri in their non‐breeding range was related to climate conditions in breeding grounds, and detected sustained population declines between 1995 and 2012 in both subspecies (L. l. menzbieri, –6.7% and L. l. baueri, –2.1% year^–1). To investigate the possible causes of the declines, we quantified changes in habitat extent at 22 migratory staging sites in the Yellow Sea, East Asia, over a 25‐year period and found –1.7% and –1.2% year^–1 loss of habitat at staging sites used by L. l. menzbieri and L. l baueri, respectively. Our results highlight the need to identify environmental and anthropogenic drivers of population change across all stages of migration to allow the formulation of effective conservation strategies across entire migratory ranges.     

Increase in protandry over time in a long‐distance migratory bird

Protandry is a widespread life‐history phenomenon describing how males precede females at the site or state of reproduction. In migratory birds, protandry has an important influence on individual fitness, the migratory syndrome, and phenological response to climate change. Despite its significance, accurate analyses on the dynamics of protandry using data sets collected at the breeding site, are lacking. Basing our study on records collected during two time periods, 1979 to 1988 and 2006 to 2016, we aim to investigate protandry dynamics over 38 years in a breeding population of willow warblers (Phylloscopus trochilus). Change in the timing of arrival was analyzed in males and females, and protandry (number of days between male and female arrival) was investigated both at population level and within breeding pairs. Our results show advancement in the arrival time at the breeding site in both sexes, but male arrival has advanced to a greater extent, leading to an increase in protandry both at the population level and within breeding pairs. We did not observe any change in sex ratio that could explain the protandry increase, but pronounced temperature change has occurred and been reported in the breeding area and along the migratory route. Typically, natural selection opposes too early arrival in males, but given warmer springs, this counteracting force may be relaxing, enabling an increase in protandry. We discuss whether our results suggest that climate change has induced sex‐specific effects, if these could be evolutionary and whether the timing of important life‐history stages such as arrival at the breeding site may change at different rates in males and females following environmental shifts.     

Longitudinal data reveal ontogenetic changes in the wing morphology of a long‐distance migratory bird

In migratory bird species, juveniles normally have shorter and more rounded wings than adults. The causes of this age‐specific difference in wing morphology, however, are largely unknown. Here, we used longitudinal data collected over 3 years from a Pied Flycatcher Ficedula hypoleuca population to assess whether age‐related differences in wing morphology are a consequence of ontogenetic changes or of selection favouring birds with longer and more pointed wings. Our study provides evidence of ontogenetic changes in wing length and shape, whereby birds grow longer and more pointed wings as they grow older. Age‐dependent variation is likely to be adaptive and may partly explain age differences in spring migration phenology and breeding success.     

Early arrival is not associated with more extra‐pair fertilizations in a long‐distance migratory bird

When assessing the benefits of early arrival date of migratory birds, a hidden and often ignored component of males’ fitness is the higher chance of early‐arriving birds to obtain extra‐pair fertilizations. Here we investigated how extra‐pair paternity might affect the relationship between male arrival date and number of fertilizations in a model study system, the European pied flycatcher Ficedula hypoleuca. For this purpose, we sampled and genotyped breeding pairs, unpaired males and offspring (including embryos from unhatched eggs when possible) of a Dutch pied flycatcher population. Detailed information on arrival date of males, egg laying date of their social mates and nest success was also recorded. Early‐arriving males had early‐laying females and males with early‐laying females had a higher probability of siring extra‐pair eggs and obtain more fertilizations. However, male arrival date alone did not correlate with the probability to gain extra‐pair paternity and neither to the amount of fertilized eggs. Both early‐ and late‐arriving males had a higher probability of losing paternity in their own nest compared to birds with an intermediate arrival date. Finally, late‐arriving males were more likely to remain unpaired but, interestingly, a few of these birds obtained paternity via extra‐pair copulations. Because earlier arrival date did not lead to more extra‐pair fertilizations and because such relationship seems to be driven mainly by the female's laying date, we conclude that the contribution of extra‐pair paternity to the overall fitness benefits of early male arrival date is relatively small.     

Ecological mismatches are moderated by local conditions for two populations of a long‐distance migratory bird

Ecological mismatches between reproductive events and seasonal resource peaks are frequently proposed to be a key driver of population dynamics resulting from global climate change. Many local populations are experiencing reduced reproductive success as a consequence of mismatches, but few mismatches have led to species‐level population declines. To better understand this apparent paradox, we investigated the breeding phenology and chick survival of two disjunct populations of Hudsonian godwits Limosa haemastica breeding at Churchill, Manitoba and Beluga River, Alaska. Only one population experienced a mismatch: godwits bred nearly one week after the onset of the invertebrate peak at Churchill because of asynchronous climatic change occurring throughout their annual cycle. However, chicks were not uniformly affected by the mismatch — growth rates and survival of young chicks were not correlated with invertebrate abundance, but older chicks tended to suffer lower survival rates on days of low invertebrate abundance. Ecological mismatches thus resulted in a complex array of consequences, but nonetheless contributed to reductions in chick survival. In contrast, godwits at Beluga River hatched their chicks just before the invertebrate peak, such that the period of highest energetic need coincided with the period of highest invertebrate abundance. As a result, growth rates and survival of godwit chicks were unaffected by invertebrate abundance. Godwits at Beluga River were able to properly time their reproduction because of predictable rates of climatic change and strong selection imposed by high predation on late‐hatched chicks. Taken together, our results suggest that population‐specific, local‐scale selection pressures play a critical role in determining the degree and severity of ecological mismatches. The potential for global climate change to induce species‐level population declines may therefore be mediated by the spatial variation in the selection pressures acting across a species’ range.     

Migration phenology and winter habitat quality are related to circulating androgen in a long‐distance migratory bird

In migratory birds, the timing of departure from wintering grounds is often dependant on the quality of habitat on an individual's territory and may influence individual fitness, resulting in an interaction of life history stages across large geographical distances. American redstart Setophaga ruticilla males who overwinter in high quality habitats arrive early to breed and subsequently produce more offspring than late arrivers. Since many migratory species overlap vernal migration with the physiological transition to breeding, we examined if breeding preparation plays a role in this seasonal interaction. We tested the hypothesis that early arriving male redstarts from high quality winter habitats are in superior breeding condition by simultaneously measuring winter habitat quality (stable‐carbon isotopes) and breeding preparation (circulating androgen, cloacal protuberance (CP) diameter) upon arrival at breeding grounds. Compared with late arrivers, early arriving males were from higher quality winter habitats and had higher androgen, but smaller CPs. Males arriving with higher androgen were in more advanced physiological migratory condition, as measured by haematocrit. Early arrivers were more likely to successfully breed, but there was no significant relationship between androgen upon arrival and breeding success. One possible explanation for these relationships is that androgen measured during arrival is most relevant in a migratory context, such that birds with high androgen may benefit from effects on migratory condition, positively influencing fitness through earlier arrival.     

“Closer‐to‐home” strategy benefits juvenile survival in a long‐distance migratory bird

Human‐induced changes in the climate and environment that occur at an unprecedented speed are challenging the existence of migratory species. Faced with these new challenges, species with diverse and flexible migratory behaviors may suffer less from population decline, as they may be better at responding to these changes by altering their migratory behavior. At the individual level, variations in migratory behavior may lead to differences in fitness and subsequently influence the population's demographic dynamics. Using lifetime GPS bio‐logging data from 169 white storks (Ciconia ciconia), we explore whether the recently shortened migration distance of storks affects their survival during different stages of their juvenile life. We also explore how other variations in migratory decisions (i.e., time, destination), movement activity (measured using overall body dynamic acceleration), and early life conditions influence juvenile survival. We observed that their first autumn migration was the riskiest period for juvenile white storks. Individuals that migrated shorter distances and fledged earlier experienced lower mortality risks. In addition, higher movement activity and overwintering “closer‐to‐home” (with 84.21% of the tracked individuals stayed Europe or North Africa) were associated with higher survival. Our study shows how avian migrants can change life history decisions over only a few decades, and thus it helps us to understand and predict how migrants respond to the rapidly changing world.     

Individual variation in orientation promotes a 3000‐km latitudinal change in wintering grounds in a long‐distance migratory raptor

Migrating juvenile birds rely on endogenous information in choosing the direction in which to fly, but such input may be overridden by social interactions with experienced individuals. We tagged seven juvenile Short‐toed Eagles Circaetus gallicus with GPS transmitters in southern Italy. This trans‐Saharan migrant flies mainly by soaring and is therefore not well adapted to performing long water crossings. Five of the seven tagged juveniles used the longer but apparently safer route towards the Strait of Gibraltar, and two migrated along a southerly trajectory and subsequently spent the winter in Sicily, apparently forced to do so by the 150‐km‐wide Sicily Channel. One of these individuals took the longer route the following autumn. These results, combined with long‐term (15 years) visual field observations involving thousands of individuals, suggest that inexperienced Short‐toed Eagles may learn their migratory routes from experienced adults, whereas some of them migrate south in response to an innate orientation instinct. Transport costs, inherited information and geography apparently interact, forcing some Short‐toed Eagles to winter 3000 km to the north of the majority of their conspecifics.     

Large‐scale,long‐term trends in British river macroinvertebrates

Rivers are among the world's most modified ecosystems, with poor water quality representing a prominent problem for over 200 years, especially in urban areas. In Western Europe, however, industrial decline, tighter regulation and improved wastewater treatment have combined over recent decades to create conditions conducive to extensive restoration and positive biological change. Here, we evaluate the river macroinvertebrate fauna of England and Wales in relation to water quality, physical habitat and climate over almost two decades. We predicted that biological recovery would be characterized by: (i) greater taxon richness and prevalence of pollution‐sensitive taxa, (ii) larger changes in more heavily urbanized catchments, and (iii) temporal trends in assemblage structure that correlated with improving water quality. Family level richness increased on average by nearly 20% during 1991–2008, accompanied by a widespread shift towards taxa characteristic of well‐oxygenated and less polluted waters. Changes were largest in the most urbanized catchments. A combination of natural gradients and anthropogenic pressures explained the variation among sites, whereas temporal changes correlated with improving water quality and variations in discharge. Positive trends were not universal, however, and there was localized deterioration in some streams draining upland areas and in the lowland south east. Our results are consistent with a large‐scale ecological recovery of English and Welsh rivers since 1990, probably continuing a trend from the mid‐20th century. Based on these results, we suggest: (i) freshwater communities are resilient to long‐term anthropogenic pressures, (ii) biodiversity benefits can arise from investment and long‐term restoration intended largely to enhance ecosystem services such as drinking water and sanitary concerns, and (iii) long‐term monitoring data collected for statutory purposes–based in this case on nearly 50 000 samples–can address scientific questions at spatial and temporal extents seldom achieved in research programmes.     

High juvenile mortality during migration in a declining population of a long‐distance migratory raptor

Many populations of long‐distance migrants are declining and there is increasing evidence that declines may be caused by factors operating outside the breeding season. Among the four vulture species breeding in the western Palaearctic, the species showing the steepest population decline, the Egyptian Vulture Neophron percnopterus, is a long‐distance migrant wintering in Africa. However, the flyways and wintering areas of the species are only known for some populations, and without knowledge of where mortality occurs, effective conservation management is not possible. We tracked 19 juvenile Egyptian Vultures from the declining breeding population on the Balkan Peninsula between 2010 and 2014 to estimate survival and identify important migratory routes and wintering areas for this species. Mortality during the first autumn migration was high (monthly survival probability 0.75) but mortality during migration was exclusively associated with suboptimal navigation. All birds from western breeding areas and three birds from central and eastern breeding areas attempted to fly south over the Mediterranean Sea, but only one in 10 birds survived this route, probably due to stronger tailwind. All eight birds using the migratory route via Turkey and the Middle East successfully completed their first autumn migration. Of 14 individual and environmental variables examined to explain why juvenile birds did or did not successfully complete their first migration, the natal origin of the bird was the most influential. We speculate that in a declining population with fewer experienced adults, an increasing proportion of juvenile birds are forced to migrate without conspecific guidance, leading to high mortality as a consequence of following sub‐optimal migratory routes. Juvenile Egyptian Vultures wintered across a vast range of the Sahel and eastern Africa, and had large movement ranges with core use areas at intermediate elevations in savannah, cropland or desert. Two birds were shot in Africa, where several significant threats exist for vultures at continental scales. Given the broad distribution of the birds and threats, effective conservation in Africa will be challenging and will require long‐term investment. We recommend that in the short term, more efficient conservation could target narrow migration corridors in southern Turkey and the Middle East, and known congregation sites in African wintering areas.     

Large‐scale,long‐term ecosystem monitoring: Interview with David Lindenmayer

David Lindenmayer leads six large‐scale, long‐term research programmes in south‐eastern Australia to help conserve biodiversity in restored areas on farmland as well as conservation in reserves, national parks, wood production forests and plantations. What makes this award‐winning researcher tick and how is the research influencing biodiversity management and policy?     

Adaptive fault bar distribution in a long‐distance migratory,aerial forager passerine?

Fault bars are translucent bands produced by stressful events during feather formation. They weaken feathers and increase their probability of breakage, and thus could compromise bird fitness by lowering flight performance. It has been recently suggested ('fault bar allocation hypothesis') that birds could have evolved adaptive mechanisms for reducing fault bar load on the feathers with the highest function during flight. We tested this hypothesis by studying first-year individuals of the long-distance migratory, aerial forager barn swallow Hirundo rustica . We predicted that fault bars should be less abundant on the outermost wing and tail feathers, but more frequent on the tail than on the outermost wing feathers. Accordingly, we found that fault bars occurred more often in tertials than in primaries or secondaries. Tail feathers had fewer fault bars than tertials, but more than primaries. Within the tail, the distribution pattern of fault bars was W-shaped, with the highest fault bar load occurring on the streamers and on the two central feathers. Because streamers are the most important tail feathers for flight performance, this finding seems to contradict the 'fault bar allocation hypothesis'. However, flight performance is much less sensitive to changes in the shape of the tail than of the wings, which could explain why evolutionary forces have not counteracted the increase of fault bars associated with feather elongation during the recent evolution of streamers in the tail of hirundines.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 455–461.     

Body size shapes inter‐specific migratory behaviour: evidence from individual tracks of long‐distance migratory shorebirds

Migration is a common phenomenon across many animal taxa. Understanding how migration scales with body size across species is fundamental in the development of migration theory and in making size‐related predictions. Although aerodynamic theory and ecophysiological scaling laws have assisted greatly in generating such predictions, their verifications have been limited by a lack of empirical data across a range of body sizes. The recent development of ultra‐light tracking devices and its rapid application to migration now allows us to put theory to the test. We used tracking data of seven closely related migratory sandpiper species (family Scolopacidae) along the East Asian‐Australasian Flyway to compare their migratory behaviour when migrating towards the breeding grounds as a function of size (50–750 g). We found a marked decline in migration speed (migration distance divided by total migration duration, including time at stopover sites and in flight) with size. Departure date from the non‐breeding (i.e. wintering) ground and arrival date at the breeding ground also scaled negatively with size. Total migration duration, migration distance, total staging duration (the number of days staying at stopover sites plus days preparing, i.e. fuelling, prior to initial migration) and step length (distance covered within one migratory leg) were not significantly related with size. Correction for phylogeny showed consistent results for all variables. Besides improving our fundamental understanding of inter‐specific variation in migration behaviours, the finding of a clear scaling with size in migration speed and migration timing highlight differential size related capabilities and constraints of migrants. Migratory birds, including sandpipers, are declining on a global scale and particularly along the East Asian‐Australasian Flyway. This notion of size‐dependency in migratory traits may have a bearing on their vulnerability to specific environmental disturbances along their flyways.