Quantification of bird migration by radar – a detection probability problem

Besides the scientific interest in the quantification of bird migration, there is an increasing need to quantify bird movements for the assessment of bird collision risk with artificial structures. In many environmental impact studies, the radar method is used in an inappropriate manner. The processing of echoes consists often of counting blips within defined screen fields, and the surveyed volume is estimated without reference to the detection probabilities of different 'target sizes' (radar cross-sections). The aim of this paper is to present a procedure to quantify bird migration reliably using radar by stating the theoretical requirements of every single step of this procedure and presenting methodological solutions using our own radar data from extensive field studies. Our methodological solutions can be applied to various radar systems, including widely used ship radar. The procedure presented involves discriminating the echoes of birds and insects and estimating the different detection probabilities of differently 'sized' birds (radar cross-sections). By ignoring the different detection probabilities, density estimations may be wrong by as much as 400%. We fear that quantification of bird migration and predicted bird numbers affected by collisions with artificial structures are in many cases based on unreliable estimates.     

A comparison of nocturnal call counts of migrating birds and reflectivity measurements on Doppler radar

Several studies have found that the peak in bird density in the atmosphere during nocturnal migration occurs before midnight, while the peak in vocalizations from migrating birds occurs after midnight, in the hours just before dawn. In a recent study, the patterns of calling from a single species of migrating birds correlated well with the patterns of density estimates of migrating birds. We test the null hypothesis that the patterns of reflectivity measurements and number of vocalizations during nocturnal migration are not related. We sampled radar data and nocturnal flight calls during spring and fall 2000 in northwestern South Carolina and southeastern New York. We analyzed changes in the hour-to-hour patterns of bird density and vocalizations for 556 hours on 58 nights. We also analyzed the night-to-night changes in the patterns of peak hour bird density and peak hour of vocalizations on 32 nights. We found that most of the hour-to-hour and night-to-night patterns of density and vocalization counts are significantly related and reject the null hypothesis. However, despite significant relationships between reflectivity measurements and vocalization counts, a great deal of variation in vocalization counts remains unexplained. These results suggest that factors other than bird density are responsible for the variation in vocalizing by migrating birds.     

Long-distance migrating species of birds travel in larger groups

How individuals migrate over long distances is an enduring mystery of animal migration. Strong selection pressure for travelling in groups has been suggested in long-distance migrating species. Travelling in groups can reduce the energetic demands of long migration, increase navigational accuracy and favour group foraging at migratory halts. Nevertheless, this hypothesis has received scant attention. I examined evolutionary transitions in migration distance in all North American breeding species of birds. I documented 72 evolutionary shifts in migration distance in the pool of 409 species. In contrasting clades, long-distance migration, as opposed to short-distance migration, was associated with a larger travelling group size. No other transitions occurred alongside in other traits such as group size in the non-breeding season or body mass. The results suggest that larger group sizes have been beneficial in the evolution of long-distance migration in a large clade of birds.     

Extracting bird migration information from C‐band Doppler weather radars

Although radar has been used in studies of bird migration for 60 years, there is still no network in Europe for comprehensive monitoring of bird migration. Europe has a dense network of military air surveillance radars but most systems are not directly suitable for reliable bird monitoring. Since the early 1990s, Doppler radars and wind profilers have been introduced in meteorology to measure wind. These wind measurements are known to be contaminated with insect and bird echoes. The aim of the present research is to assess how bird migration information can be deduced from meteorological Doppler radar output. We compare the observations on migrating birds using a dedicated X‐band bird radar with those using a C‐band Doppler weather radar. The observations were collected in the Netherlands, from 1 March to 22 May 2003. In this period, the bird radar showed that densities of more than one bird per km³ are present in 20% of all measurements. Among these measurements, the weather radar correctly recognized 86% of the cases when birds were present; in 38% of the cases with no birds detected by the bird radar, the weather radar claimed bird presence (false positive). The comparison showed that in this study reliable altitudinal density profiles of birds cannot be obtained from the weather radar. However, when integrated over altitude, weather radar reflectivity is correlated with bird radar density. Moreover, bird flight speeds from both radars show good agreement in 78% of cases, and flight direction in 73% of cases. The usefulness of the existing network of weather radars for deducing information on bird migration offers a great opportunity for a European‐wide monitoring network of bird migration.     

Migration of red‐backed shrikes from the Iberian Peninsula: optimal or sub‐optimal detour?

The current Northern Hemisphere migration systems are believed to have arisen since the last glaciation. In many cases, birds do not migrate strait from breeding to non‐breeding areas but fly via a detour. All western European populations of red‐backed shrikes Lanius collurio are assumed to reach their southern African wintering grounds detouring via southeast Europe. Based on theoretical considerations under an optimality framework this detour is apparently optimal. Here, we use individual geolocator data on red‐backed shrikes breeding in Spain to show that these birds do indeed detour via southeast Europe en route to southern Africa where they join other European populations of red‐backed shrikes and return via a similar route in spring. Disregarding potential wind assistance, the routes taken for the tracked birds in autumn were not optimal compared to crossing the barrier directly. For spring migration the situation was quite different with the detour apparently being optimal. However, when considering potential wind assistance estimated total air distances during autumn migration were overall similar and the barrier crossing shorter along the observed routes. We conclude that considering the potential benefit of wind assistance makes the route via southeast Europe likely to be less risky in autumn. However, it cannot be ruled out that other factors, such as following a historical colonisation route could still be important.     

Calibration of visually estimated distances to migrating seabirds with radar measurements

ABSTRACT Censusing seabirds from coastal areas requires reliable estimates of bird numbers and the distances of the birds from the coastline. Logistical constraints make visual estimation of distances the only feasible method in many studies. We tested the accuracy of visually estimated offshore distances of six migratory seabird species in the Strait of Gibraltar using simultaneous measurements obtained by radar. Most birds (91%) were detected within 3 km of the coast and we truncated our calibration at this distance. We found a strong correlation between radar and visual estimates (R²= 0.83, P < 0.0001). The magnitude of errors in visual estimates was moderate and ranged from 0.08 to 0.20 for different distances and observers. Among the factors potentially affecting the accuracy of visual estimates of distance to seabird in our study were observer identity, bird species, bird behavior, and weather; the most parsimonious model in our study included observer identity as the only predictor, and no model with more than one predictor had a smaller Akaike's information criterion value. Radar can be used to help train observers and to reduce biases in visual estimates of distances by means of calibration. When no other methods are available to accurately measure distances to seabirds, visual estimates of distances, recorded by experienced observers and once calibrated with radar (or other ground‐truthing methods), may be acceptable for different species under a wide range of environmental conditions.     

Flight mode affects allometry of migration range in birds

Billions of birds migrate to exploit seasonally available resources. The ranges of migration vary greatly among species, but the underlying mechanisms are poorly understood. I hypothesise that flight mode (flapping or soaring) and body mass affect migration range through their influence on flight energetics. Here, I compiled the tracks of migratory birds (196 species, weighing 12–10 350 g) recorded by electronic tags in the last few decades. In flapping birds, migration ranges decreased with body mass, as predicted from rapidly increasing flight cost with increasing body mass. The species with higher aspect ratio and lower wing loading had larger migration ranges. In soaring birds, migration ranges were mass‐independent and larger than those of flapping birds, reflecting their low flight costs irrespective of body mass. This study demonstrates that many animal‐tracking studies are now available to explore the general patterns and the underlying mechanisms of animal migration.     

Causes and characteristics of reverse bird migration: an analysis based on radar,radio tracking and ringing at Falsterbo,Sweden

That birds migrate in the reverse direction of the expected is a phenomenon of regular occurrence which has been observed at many sites. Here we use three different methods; tracking radar, radiotelemetry and ringing, to characterize the flights of these reverse migrants and investigate possible causes of reverse migration of nocturnally migrating passerines during autumn migration at Falsterbo peninsula, Sweden. Using these different methods we investigated both internal factors, such as age and fuel load, and external factors such as weather variables, competition and predation risk. Birds flying in the reverse direction were more likely to be lean and to be juveniles. Reverse migration was also more common with overcast skies and winds with north and east components. We did not find any effect of temperature, visibility, number of migrating sparrowhawks, or the total number of ringed birds at the site on the day of departure. We found that reverse migration is characterized by slower flight speeds (airspeed) at high altitudes and that it takes place later in the night than forward migration.     

The influence of weather on the flight altitude of nocturnal migrants in mid‐latitudes

By altering its flight altitude, a bird can change the atmospheric conditions it experiences during migration. Although many factors may influence a bird's choice of altitude, wind is generally accepted as being the most influential. However, the influence of wind is not clearly understood, particularly outside the trade‐wind zone, and other factors may play a role. We used operational weather radar to measure the flight altitudes of nocturnally migrating birds during spring and autumn in the Netherlands. We first assessed whether the nocturnal altitudinal distribution of proportional bird density could be explained by the vertical distribution of wind support using three different methods. We then used generalized additive models to assess which atmospheric variables, in addition to altitude, best explained variability in proportional bird density per altitudinal layer each night. Migrants generally remained at low altitudes, and flight altitude explained 52 and 73% of the observed variability in proportional bird density in spring and autumn, respectively. Overall, there were weak correlations between altitudinal distributions of wind support and proportional bird density. Improving tailwind support with height increased the probability of birds climbing to higher altitude, but when birds did fly higher than normal, they generally concentrated around the lowest altitude with acceptable wind conditions. The generalized additive model analysis also indicated an influence of temperature on flight altitudes, suggesting that birds avoided colder layers. These findings suggested that birds increased flight altitudes to seek out more supportive winds when wind conditions near the surface were prohibitive. Thus, birds did not select flight altitudes only to optimize wind support. Rather, they preferred to fly at low altitudes unless wind conditions there were unsupportive of migration. Overall, flight altitudes of birds in relation to environmental conditions appear to reflect a balance between different adaptive pressures.     

Artificial light at night confounds broad‐scale habitat use by migrating birds

With many of the world's migratory bird populations in alarming decline, broad‐scale assessments of responses to migratory hazards may prove crucial to successful conservation efforts. Most birds migrate at night through increasingly light‐polluted skies. Bright light sources can attract airborne migrants and lead to collisions with structures, but might also influence selection of migratory stopover habitat and thereby acquisition of food resources. We demonstrate, using multi‐year weather radar measurements of nocturnal migrants across the northeastern U.S., that autumnal migrant stopover density increased at regional scales with proximity to the brightest areas, but decreased within a few kilometers of brightly‐lit sources. This finding implies broad‐scale attraction to artificial light while airborne, impeding selection for extensive forest habitat. Given that high‐quality stopover habitat is critical to successful migration, and hindrances during migration can decrease fitness, artificial lights present a potentially heightened conservation concern for migratory bird populations.     

Why migrate during the day: a comparative analysis of North American birds

Migration can take place primarily during the day or at night, or during both depending on the species. Why the timing of migration varies among species has been the object of much research but the underlying ecological processes are still unclear. Proximally, migration timing may be influenced by the timing of other activities or may be more prevalent in species that migrate over long distances. Adaptive scenarios emphasize the reduction in flight costs at night especially in smaller species and the advantages of travelling in groups during the day to locate staging sites more efficiently. I used phylogenetic independent contrasts to examine these hypotheses in all North American nesting birds. I uncovered 24 evolutionary transitions in migration timing, most of which involved a switch from nocturnal to diurnal migration. Few of these transitions involved a concomitant change in the timing of foraging habits or migration distance. However, species in diurnal clades were larger, travelled in larger flocks and were generally more sociable than their nocturnal counterparts. The results support the hypotheses that a reduction in flight costs and the ability to pool information from companions are associated with migration timing in North American bird species.     

The natural link between Europe and Africa – 2.1 billion birds on migration

The Palaearctic–African migration system comprises enormous numbers of birds travelling between Europe and Africa twice each year. Migratory birds may form strong links between the two continents given they can act as both transport vehicles for parasites and diseases as well as temporary consumers with increased food demand to fuel their flight. Knowing the number of migrating birds is crucial if such links are to be quantified. We estimate that today approximately 2.1 billion songbirds and near-passerine birds migrate from Europe to Africa in autumn, 73% of which are accounted for by just 16 species. This number is only half the estimate from the 1950s in the only other assessment to date. The discrepancy is mainly caused by the limited information on population sizes in the past. Our estimated number of migrants is highly dependent on the accuracy of the underlying estimates of breeding population sizes, as well as breeding parameters in species with relatively high reproductive output. The updated figures quantify and emphasize the strong natural connection between Africa and Europe, which has important implications for manifold research topics including those related to climate change, human health and biological conservation.     

Young and female‐biased irruptions in pygmy owls Glaucidium passerinum in southern Finland

Irruptive migrants often show biased sex and age ratio, and typically young females are the most abundant class participating in irruptions. Several hypotheses have been proposed to explain differential migration in birds. We examined these hypotheses in a sexually size‐dimorphic species (females being larger than males), the pygmy owl Glaucidium passerinum, whose migration behaviour has also been poorly documented. Migration data were collected at the Hanko Bird Observatory, SW Finland during 1979–2010. Pygmy owls showed large fluctuations in migration numbers between years. Most of the migrants were young birds and females migrated in larger numbers than males. Results support the ‘territory defence hypothesis’ since males showed weaker tendency to migrate despite the smaller body size, but the ‘dominance hypothesis’ was also supported (subordinate young outnumber dominant adults in migration numbers). Findings support the idea that in owl species, which are dependent on nesting cavities, males show lower migration behaviour than females, whereas the pattern is opposite in species which normally breed outside cavities. In addition, adults migrated later than young, likely because they need to moult remiges before departure. Early hatched young also migrated earlier than late hatched young, suggesting that more dominant young migrate first.     

Genetic divergence and migration patterns in a North American passerine bird: implications for evolution and conservation

Like many other migratory birds, the black-throated blue warbler (Dendroica caerulescens) shows pronounced differences in migratory behaviour and other traits between populations: birds in the southern part of the breeding range have darker plumage and migrate to the eastern Caribbean during the winter, whereas those in the north have lighter plumage and migrate to the western Caribbean. We examined the phylogeography of this species, using samples collected from northern and southern populations, to determine whether differentiation between these populations dates to the Pleistocene or earlier, or whether differences in plumage and migratory behaviour have arisen more recently. We analysed variation at 369 bp of the mitochondrial control region domain I and also at seven nuclear microsatellites. Analyses revealed considerable genetic variation, but the vast majority of this variation was found within rather than between populations, and there was little differentiation between northern and southern populations. Phylogeographic analyses revealed a very shallow phylogenetic tree, a star-like haplotype network, and a unimodal mismatch distribution, all indicative of a recent range expansion from a single refugium. Coalescent modelling approaches also indicated a recent common ancestor for the entire group of birds analysed, no split between northern and southern populations, and high levels of gene flow. These results show that Pleistocene or earlier events have played little role in creating differences between northern and southern populations, suggesting that migratory and other differences between populations have arisen very recently. The implications of these results for the evolution of migration and defining taxonomic groups for conservation efforts are discussed.     

鸟类迁徙:在全球变暖趋势下的演化、调控与发展(英文)

最近几十年的研究证实 ,鸟类迁徙在很大程度上受到遗传因素的直接控制。有证据表明 ,存在某种先天的迁徙动因并涉及以下几方面的遗传调控 :(1)迁徙过程的起始、持续以及结束 ;(2 )迁徙活动量 ,即决定鸟类飞行距离的遗传参数 ;(3)迁徙方向 ;(4)生理参数 ,特别是迁徙期间的脂肪贮存 ,以及对于那些部分个体迁徙的鸟种而言 ,决定个体迁徙与否的生理参数。双因素选择实验表明 ,部分迁徙群经由几个世代的选择即可转变成完全的迁徙群或非迁徙群。新迁徙方向以及由此导致的新越冬区的改变 ,也能在野生鸟类中迅速实现。至少在以往研究得最为透彻的鸟种 (黑顶林莺Sylviaatricapilla)中 ,“迁徙”或“非迁徙”是先天性的 ,与特异性迁徙活动量相关 (尤如一时间程序 ) ,前者 (迁徙的 )已证实是由一种阈机制所控制的。一项新的鸟类迁徙理论假设 ,即使好些完全迁徙的类群 ,较低水平的迁徙活动量选择也会导致阈的异位 ,低于这一阈值就会出现非迁徙个体。因此 ,通过选择作用 ,一个迁徙型种群可以通过部分迁徙型转变为非迁徙型。这种中间阶段在现存鸟类中十分普遍。它始见于生物演化早期 ,就鸟类而言 ,可能在原始鸟类就已具备。模型运算表明 ,在施以强定向选择情况下 ,迁徙鸟类经过约 4 0年可转变为留鸟 ,反之亦然。这就解     

ROOSTING MOVEMENTS OF BIRDS AND MIGRATION DEPARTURES FROM ROOSTS AS SEEN BY RADAR

Unusual radar echoes attributable to the roosting movements of large flocks of birds, and to departures from roosts on migration flight are illustrated. Probably the most striking displays are received from Starlings Sturnus vulgaris. Radar evidence is shown of a much more rapid break-up of flocks on migration at night than by day. Weather at departure points is studied.     

Heterothermy,body size,and locomotion as ecological predictors of migration in mammals

1. Migration is ubiquitous among animals and has evolved repeatedly and independently. Comparative studies of the evolutionary origins of migration in birds are widespread, but are lacking in mammals. Mammalian species have greater variation in functional traits that may be relevant for migration. Interspecific variation in migration behaviour is often attributed to mode of locomotion (i.e. running, swimming, and flying) and body size, but traits associated with the evolutionary precursor hypothesis, including geographic distribution, habitat, and diet, could also be important predictors of migration in mammals. Furthermore, mammals vary in thermoregulatory strategies and include many heterothermic species, providing an alternative strategy to avoid seasonal resource depletion.
2. We tested the evolutionary precursor hypothesis for the evolution of migration in mammals and tested predictions linking migration to locomotion, body size, geographic distribution, habitat, diet, and thermoregulation. We compiled a dataset of 722 species from 27 mammalian orders and conducted a series of analyses using phylogenetically informed models.
3. Swimming and flying mammals were more likely to migrate than running mammals, and larger species were more likely to migrate than smaller ones. However, heterothermy was common among small running mammals that were unlikely to migrate. High-latitude swimming and flying mammals were more likely to migrate than high-latitude running mammals (where heterothermy was common), and most migratory running mammals were herbivorous. Running mammals and frugivorous bats with high thermoregulatory scope (greater capacity for heterothermy) were less likely to migrate, while insectivorous bats with high thermoregulatory scope were more likely to migrate.
4. Our results indicate a broad range of factors that influence migration, depending on locomotion, body size, and thermoregulation. Our analysis of migration in mammals provided insight into some of the general rules of migration, and we highlight opportunities for future investigations of exceptions to these rules, ultimately leading to a comprehensive understanding of the evolution of migration.

     

Comparing transect survey and WSR‐88D radar methods for monitoring daily changes in stopover migrant communities

ABSTRACT For decades, researchers have successfully used ground‐based surveys to understand localized spatial and temporal patterns in stopover habitat use by migratory birds. Recent technological advances with WSR‐88D radar now allow such investigations on much broader spatial scales. Both methods are assumed to accurately quantify patterns in migrant bird communities, yet information is lacking regarding relationships between radar estimates of migration and different ground‐based monitoring methods. From 2005 to 2007, we monitored migrant communities on or near two Department of Defense installations in the spring (Ft. Polk Military Complex, LA; U.S. Army Test and Evaluation Command, Yuma Proving Ground, AZ) and on two installations in the fall (Ft. Polk Military Complex, LA; Eglin Air Force Base, FL) using both ground‐based transect surveys and radar imagery of birds aloft. We modeled daily changes in migrant abundance and positive and negative species turnover measured on the ground as a function of radar estimates of migrant exodus and input densities. Radar data were not significant predictors of any response variable in any season either in the southeastern or southwestern United States, indicating a disparity between the results obtained using different methods. Multiple unique sources of error associated with each technique likely contributed to the conflicting outcomes, and researchers should take great care when selecting monitoring methods appropriate to address research questions, effects of management practices, or when comparing the results of migration studies using different survey techniques.     

Farmland as stopover habitat for migrating birds – effects of organic farming and landscape structure

Agricultural intensification in Europe has affected farmland bird populations negatively, both during summer and winter. Although the migratory period poses separate challenges on birds than breeding and wintering, the consequences of farming practices for birds during migration remain poorly investigated. We monitored abundance and species richness of migratory birds in autumn at matched pairs of organic and conventional farms situated either in intensively farmed open plains (homogeneous landscapes) or in small‐scale farming landscapes (heterogeneous landscapes) in southern Sweden. Total bird density did not differ between landscape types but was marginally higher on organic compared to conventional farms. When including taxonomic status in the model (passerines vs non‐passerines), we found significantly more birds on organic farms, and more non‐passerines in the homogeneous landscapes. The effect of farming practice and landscape type on density differed between functional groups. Omnivore density was higher in the homogeneous landscapes, and invertebrate feeders were marginally more abundant on organic farms. The effects of farming practice on the overall species richness and on the density of granivorous birds were landscape dependent. In the homogeneous landscapes, organic farms held a higher number of species and density of granivorous birds than conventional farms, but there was no such difference in the heterogeneous landscapes. Thus, organic farming can enhance abundance and species richness of farmland birds during migration, but the effect differs between landscape types and species. The effectiveness of organic farming was highest in the homogeneous landscape making it important to promote organic farming there. However, for some species during migration, increased heterogeneity in homogeneous landscapes may have negative effects. We propose that migratory bird diversity in homogeneous landscapes may be best preserved by keeping the landscape open, but that a reduced agricultural intensity, such as organic farming, should be encouraged.     

THE MIGRATION PATTERNS OF THE PURPLE SANDPIPER CALIDRIS MARITIMA

Summers, R. W. 1994. The migration patterns of the Purple Sandpiper Calidris maritima. Ostrich 65: 167–173.

The Purple Sandpiper breeds largely in the Arctic, and winters (boreal season) on the rocky shores of the north Atlantic, further north than any other sandpiper. As the populations from Canada, Greenland, Iceland, Svalbard, Norway and Russia differ in wing and bill lengths it is possible to match measurements taken from breeding birds with samples of birds caught in winter. Ringing recoveries, especially from colour marked birds, have also helped to determine migration routes and wintering areas. Four populations move to the nearest ice-free coast. Two populations move south of the nearest ice-free coast, being replaced by larger birds from a more northerly population (“chain migration”). Only the north Canadian population is believed to migrate a long distance, “leap-frogging” other winter populations. These patterns are discussed in relation to theories for the migration patterns of waders.