Flight by night or day? Optimal daily timing of bird migration

Many migratory bird species fly mainly during the night (nocturnal migrants), others during daytime (diurnal migrants) and still others during both night and day. Need to forage during the day, atmospheric structure, predator avoidance and orientation conditions have been proposed as explanations for the widespread occurrence of nocturnal migration. However, the general principles that determine the basic nocturnal-diurnal variation in flight habits are poorly known. In the present study optimal timing of migratory flights, giving the minimum total duration of the migratory journey, is evaluated in a schematic way in relation to ecological conditions for energy gain in foraging and for energy costs in flight. There exists a strong and fundamental advantage of flying by night because foraging time is maximized and energy deposition can take place on days immediately after and prior to the nocturnal flights. The increase in migration speed by nocturnal compared with diurnal migration will be largest for birds with low flight costs and high energy deposition rates. Diurnal migration will be optimal if it is associated with efficient energy gain immediately after a migratory flight because suitable stopover/foraging places have been located during the flight or if energy losses during flight are substantially reduced by thermal soaring and/or by fly-and-forage migration. A strategy of combined diurnal and nocturnal migration may be optimal when birds migrate across regions with relatively poor conditions for energy deposition (not only severe but also soft barriers). Predictions about variable timing of migratory flights depending on changing foraging and environmental conditions along the migration route may be tested for individual birds by analysing satellite tracking results with respect to daily travel routines in different regions. Documenting and understanding the adaptive variability in daily travel schedules among migrating animals constitute a fascinating challenge for future research.     

Effects of weather on avian migration at proposed ridgeline wind energy sites

With the popularity of wind energy increasing globally, concerns surfaced in the 1980s as to the potential adverse effects of wind turbines on migrating birds. Understanding how weather conditions influence passage rates can help determine the potential for increased avian–turbine collisions. Using vertical and horizontal mounted marine radars, raptor stand watch observations, and portable handheld weather stations, we studied how temperature, cloud cover, barometric pressure, wind direction, and wind speed affected avian passage rates and height of migrants over 3 ridges (Wartenbe, North Dokie, and South Dokie) being developed for wind energy in northern British Columbia. Using an Akaike's Information Criterion (AIC), we determined that a reduced model combining wind speed, barometric pressure, and cloud cover was best at explaining and predicting higher passage rates (expressed as no. birds/hr) in the fall migration for both diurnal and nocturnal migrants. Wind speed proved the most important predictor of passage rates for spring nocturnal migrants and a combination of cloud cover, temperature, and wind direction for diurnal spring migrants. Wind speed also predicted decreases in flight altitude among nocturnal migrants but increased altitude in diurnal migrants. This information coupled with migration timing and topographical areas of higher migrant activity can be useful to wind energy proponents who wish to mitigate collision risk with migrating birds. © 2011 The Wildlife Society.     

Revealing patterns of nocturnal migration using the European weather radar network

Nocturnal avian migration flyways remain an elusive concept, as we have largely lacked methods to map their full extent. We used the network of European weather radars to investigate nocturnal bird movements at the scale of the European flyway. We mapped the main migration directions and showed the intensity of movement across part of Europe by extracting biological information from 70 weather radar stations from northern Scandinavia to Portugal, during the autumn migration season of 2016. On average, over the 20 nights and all sites, 389 birds passed per 1 km transect per hour. The night with highest migration intensity showed an average of 1621 birds km^–1 h^–1 passing the radar stations, but there was considerable geographical and temporal variation in migration intensity. The highest intensity of migration was seen in central France. The overall migration directions showed strong southwest components. Migration dynamics were strongly related to synoptic wind conditions. A wind‐related mass migration event occurred immediately after a change in wind conditions, but quickly diminished even when supporting winds continued to prevail. This first continental‐scale study using the European network of weather radars demonstrates the wealth of information available and its potential for investigating large‐scale bird movements, with consequences for ecosystem function, nutrient transfer, human and livestock health, and civil and military aviation.     

THE NIGHTLY INITIATION OF PASSERINE MIGRATION IN SPRING: A DIRECT VISUAL STUDY

The nightly initiation of migration of passerine birds was studied during a spring season in coastal Louisiana. A horizontally-directed portable ceilometer placed on a tower illuminated birds as they took off from or landed in the trees. Daily censuses were conducted to supplement nocturnal observations. Local weather conditions were continuously monitored at the site of the study. Appreciable numbers of migrants landed in the woods on the coast almost every day, the first birds usually appearing between 10.30 and 11.00 hrs, and the last about 16.00 hrs. On two occasions birds were seen landing after dark. The migrants usually became quiet by about 18.00 hrs and the first bird was usually seen leaving the trees just over an hour later. The exodus usually peaked between 19.10 and 19.15 hrs, 40 to 45 minutes after sunset. The time of initiation of nocturnal migration was not significantly affected by immediate weather factors. The duration of the exodus varied widely, and was related to the number of individual birds in the census area. On some occasions grounded migrants stayed overnight, but only when there had been an active cold front over the Gulf. No local weather conditions examined were found to be directly inhibitory to nocturnal migration. Birds departed singly and without preliminary activity. Wind direction influenced the initial orientation of the migrants. With southerly winds, 57% of the birds seen flying across the beam initially headed toward the Gulf, while with northerly winds only 19% initially flew in this direction.     

Nocturnal migrants do not incur higher collision risk at wind turbines than diurnally active species

Nocturnally migrating birds, particularly passerines, are known to be vulnerable to collision with man‐made structures such as buildings, towers or offshore platforms, yet information with respect to wind farms is ambiguous. We recorded bird flight intensities using radar during autumn migration at four wind farms situated within a major migration flyway in northern Germany and simultaneously conducted systematic searches for collision fatalities at the same sites. We found that migration traffic rates at rotor height estimated by radar observations were significantly higher during the night, yet strictly nocturnal migrants constituted only 8.6% of all fatalities at the wind farms. In contrast to the situation at other vertical structures, nocturnal migrants do not have a higher risk of collision with wind energy facilities than do diurnally active species, but rather appear to circumvent collision more effectively.     

Timing of nocturnal autumn migratory departures in juvenile European robins (Erithacus rubecula) and endogenous and external factors

Many passerine medium distance nocturnal migrants take off from stopover sites not only at the beginning of the night, but also in the middle and at the end of the night. In this paper, we tested two explanations for this phenomenon: (1) that departure time is governed by fuel stores, and (2) that departure time is influenced by the weather. The relationship of temporal distribution of migratory nocturnal departures with body condition and weather factors was studied in juvenile European robins (Erithacus rubecula) during autumn migration. The study was done on the Courish Spit on the Baltic Sea in 1997–2003 by retrapping 74 ringed birds in high mist nets during nocturnal migratory departure. Departure time was not related to fuel stores at arrival and departure, stopover duration, fuel deposition rate or progress of the season. Nor did the local weather at departure influence departure time. A possible reason was a large variation in the behaviour of the birds. European robins which made 1-day stopovers arrived and departed during better weather conditions than birds that stopped over for longer periods. In the former cohort, a significant model with four predictors explained 55% of variation in departure time. It is assumed that weather at the night of departure and during the previous night influenced the time of take-offs in these birds. In robins which made long stopovers, departure time is probably governed by their individual endogenous circadian rhythms of activity, which are related to the environment in a complex way.     

Bird movements at rotor heights measured continuously with vertical radar at a Dutch offshore wind farm

Assessing the impacts of avian collisions with wind turbines requires reliable estimates of avian flight intensities and altitudes, to enable accurate estimation of collision rates, avoidance rates and related effects on populations. At sea, obtaining such estimates visually is limited not only by weather conditions but, more importantly, because a high proportion of birds fly at night and at heights above the range of visual observation. We used vertical radar with automated bird‐tracking software to overcome these limitations and obtain data on the magnitude, timing and altitude of local bird movements and seasonal migration measured continuously at a Dutch offshore wind farm. An estimated 1.6 million radar echoes representing individual birds or flocks were recorded crossing the wind farm annually at altitudes between 25 and 115 m (the rotor‐swept zone). The majority of these fluxes consisted of gull species during the day and migrating passerines at night. We demonstrate daily, monthly and seasonal patterns in fluxes at rotor heights and the influence of wind direction on flight intensity. These data are among the first to show the magnitude and variation of low‐altitude flight activity across the North Sea, and are valuable for assessing the consequences of developments such as offshore wind farms for birds.     

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.     

Monitoring flight calls of migrating birds from an oil platform in the northern Gulf of Mexico

ABSTRACT.   Millions of birds migrate across the Gulf of Mexico each year. However, most studies of migration in this region involve sampling onshore locations during the day, potentially underrepresenting the diversity and abundance of migrants passing the region. We evaluated a potential solution to this problem by recording the flight calls of passing migrants from an oil platform located southeast of the Alabama coast in the Gulf of Mexico. We detected 2762 calls during 30 nights from 9 September to 2 November 1999, and were able to identify 2329 calls to species. Flight calls by nine species of birds represented 23% of all identified calls. The greatest number of calls during one night (1017 calls) and during a 1-h period (257 calls) were recorded on 10 September. The greatest number of calls was recorded 8 h after sunset, with a secondary peak 2 h after sunset. The peak prior to sunrise may indicate the formation of flocks at dawn, and the peak after sunset may have been caused by the first wave of migrants reaching the platform. However, call counts varied extensively, with 98% of all calls recorded during 13 nights and 40% on a single night, possibly resulting from hourly and nightly differences in bird numbers aloft, atmospheric conditions, and artificial lighting conditions. Although recording on oil platforms can be difficult because of mechanical, wind, and wave noise, our results suggest great potential for describing the species composition of passing vocal migrants and the temporal patterns of flight-calling behavior if quiet recording locations can be found. Moreover, flight call monitoring could be a critically important tool for bird conservation in this region, given recent proposals to develop wind power and the potential bird mortality associated with such developments.     

Proximate Cues for a Short-Distance Migratory Species: an Application of Survival Analysis

ABSTRACT Investigation of bird migration has often highlighted the importance of external factors in determining timing of migration. However, little distinction has been made between short- and long-distance migrants and between local and flight birds (passage migrants) in describing migration chronology. In addition, measures of food abundance as a proximate factor influencing timing of migration are lacking in studies of migration chronology. To address the relationship between environmental variables and timing of migration, we quantified the relative importance of proximate external factors on migration chronology of local American woodcock (Scolopax minor), a short distance migrant, using event-time analysis methods (survival analysis). We captured 1,094 woodcock local to our study sites in Michigan, Minnesota, and Wisconsin (USA) during autumn 2002–2004 and documented 786 departure dates for these birds. Photoperiod appeared to provide an initial proximate cue for timing of departure. Moon phase was important in modifying timing of departure, which may serve as a navigational aid in piloting and possibly orientation. Local synoptic weather variables also contributed to timing of departure by changing the rate of departure from our study sites. We found no evidence that food availability influenced timing of woodcock departure. Our results suggest that woodcock use a conservative photoperiod-controlled strategy with proximate modifiers for timing of migration rather than relying on abundance of their primary food, earthworms. Managing harvest pressure on local birds by adjusting season lengths may be an effective management tool with consistent migration patterns from year to year based on photoperiod.     

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.     

A place to land: spatiotemporal drivers of stopover habitat use by migrating birds

Migrating birds require en route habitats to rest and refuel. Yet, habitat use has never been integrated with passage to understand the factors that determine where and when birds stopover during spring and autumn migration. Here, we introduce the stopover‐to‐passage ratio (SPR), the percentage of passage migrants that stop in an area, and use 8 years of data from 12 weather surveillance radars to estimate over 50% SPR during spring and autumn through the Gulf of Mexico and Atlantic coasts of the south‐eastern US, the most prominent corridor for North America’s migratory birds. During stopovers, birds concentrated close to the coast during spring and inland in forested landscapes during autumn, suggesting seasonal differences in habitat function and highlighting the vital role of stopover habitats in sustaining migratory communities. Beyond advancing understanding of migration ecology, SPR will facilitate conservation through identification of sites that are disproportionally selected for stopover by migrating birds.     

Winds at departure shape seasonal patterns of nocturnal bird migration over the North Sea

On their migratory journeys, terrestrial birds can come across large inhospitable areas with limited opportunities to rest and refuel. Flight over these areas poses a risk especially when wind conditions en route are adverse, in which case inhospitable areas can act as an ecological barrier for terrestrial migrants. Thus, within the east-Atlantic flyway, the North Sea can function as an ecological barrier. The main aim of this study was to shed light on seasonal patterns of bird migration in the southern North Sea and determine whether departure decisions on nights of intense migration were related to increased wind assistance. We measured migration characteristics with a radar that was located 18 km off the NW Dutch coast and used simulation models to infer potential departure locations of birds on nights with intense nocturnal bird migration. We calculated headings, track directions, airspeeds, groundspeeds on weak and intense migration nights in both seasons and compared speeds between seasons. Moreover, we tested if departure decisions on intense migration nights were associated with supportive winds. Our results reveal that on the intense migration nights in spring, the mean heading was towards E, and birds departed predominantly from the UK. On intense migration nights in autumn, the majority of birds departed from Denmark, Germany and north of the Netherlands with the mean heading towards SW. Prevailing winds from WSW at departure were supportive of a direct crossing of the North Sea in spring. However, in autumn winds were generally not supportive, which is why many birds exploited positive wind assistance which occurred on intense migration nights. This implies that the seasonal wind regimes over the North Sea alter its migratory dynamics which is reflected in headings, timing and intensity of migration.     

Conquering the night: understanding nocturnal migration in birds

Abstract

Migration is a biologically distinct and unique phenomenon that enables the birds to migrate twice-a-year between the breeding and wintering grounds. These movements are known as spring and autumn migration, respectively. Depending on their inherent programming, the migratory birds may fly during day or night or both. Different environmental factors such as, temperature, food, predator pressure and physiological demands of energy storage and expenditure, contribute to the pattern of migrations, day or nighttime. Since, most of them are nighttime migrants they have to make dramatic changes in their physiology and behavior to transform them from being diurnal to predominantly nocturnal. These changes result in different life history stages (LHSs) such as migration, reproduction and molt, in their annual cycle, which are regulated by endogenous circadian and circannual clocks. As a result, the birds start preparing well in advance for the approaching LHS. The present review focuses on behavioral strategies of a nocturnal migrant and understanding of the possible physiological responses to ensure successful migration.     

Within night correlations between radar and ground counts of migrating songbirds

ABSTRACT.   Studies comparing numbers of nocturnal migrants in flight with numbers of migrants at stopover sites have produced equivocal results. In 2003, we compared numbers of nocturnal migrants detected by radar to numbers of passerines observed at the Atlantic Bird Observatory in southwestern Nova Scotia, Canada. Numbers of nocturnal migrants detected by radar were positively correlated with numbers of migrants as determined by mist-netting, censuses, and daily estimated totals (daily estimates of birds present based netting and census results and casual observations) the following day. On nights with winds favorable for migration (tailwinds), the peak correlation between ground counts and radar counts the night before occurred just after sunset. On nights with unfavorable winds (headwinds), the correlation increased through the night, with a peak just before sunrise. The patterns of correlation are consistent with a scenario where birds accumulate at the coastline during periods of unfavorable wind, likely because they are not willing to cross a major ecological barrier, the Gulf of Maine. On nights with favorable winds, many birds departed, but some, possibly after testing wind conditions, apparently decided not to cross the Gulf of Maine and returned. Our results suggest that combining data collected using different methods to generate a daily estimated total provides the best estimate of the number of migrants present at a stopover site. Simultaneous studies at multiple locations where different census methods are used, making more effective use of temporal data (both from radar and diurnal counts), will more clearly elucidate patterns of flight behavior by migratory songbirds and the relationship between ground counts and counts of birds aloft.     

沈阳桃仙国际机场鸟类夜间迁徙规律

在鸟类迁徙季节,夜间鸟击事故频发是机场鸟击发生的一个显著特点.了解鸟类的夜间迁徙规律对于改进夜间鸟击防范措施具有重要的指导意义.本研究综合采用网捕法和声音记录法对沈阳桃仙机场夜间鸟类迁徙物种组成和迁徙规律进行研究.结果表明: 56种鸟类(占比88.9%)具有夜间迁徙习性,且以后半夜迁徙为主;鸟类夜间迁徙具有明显的时间动态和迁徙次序,春季鸟类迁徙较为集中,迁徙高峰在5月中旬,主要鸟类由鹌鹑、红尾伯劳、栗耳鹀、黑喉石鵖、普通夜鹰、黄眉柳莺等组成,秋季迁徙较为分散,迁徙高峰出现在9月下旬至10月上旬,主要由鹌鹑、灰背鸫、红喉鹨、丘鹬、矛斑蝗莺和灰头鵐等组成.对夜间迁徙鸟类的危险等级评估发现,春季严重危险物种是鹌鹑和红尾伯劳,秋季严重危险物种是鹌鹑、纵纹腹小鸮、灰背鸫和丘鹬.分别从夜间迁徙鸟类组成、迁徙动态、时间节律和物种危险等级等角度提出了相应的鸟击防范对策,为桃仙机场鸟击防范提供参考.     

Altitudinal migration by birds: a review of the literature and a comprehensive list of species

Altitudinal migration is the seasonal altitudinal movement of birds from breeding areas to non‐breeding or wintering areas at different elevations. Although this type of migration is widely reported, questions remain concerning the number of species that perform altitudinal migration, possible variation among different taxa and geographic locations in the extent of altitudinal migration, and the foraging guilds of altitudinal migrants. We conducted an extensive bibliographic survey and compiled a list of altitudinal migrant birds worldwide. We characterized species in terms of their foraging guilds because the spatial distribution of food resources along altitudinal gradients is often evoked as a driver of bird altitudinal migration. We identified 1238 species of altitudinal migrants, ~10% of the ~10,000 extant species of birds. We found a strong geographic bias in publications focusing on avian altitudinal migration toward the United States and Costa Rica, and a paucity of studies in megadiverse regions such as the Afrotropical and Indomalayan realms, and areas in the Neotropics other than Costa Rica. We also found that most species of altitudinal migrants were invertivores rather than frugivores or nectarivores. This general pattern held true for all zoogeographic realms except the Neotropics, where nectarivores and frugivores predominated among altitudinal migrants. The prevalence of invertivore birds among altitudinal migrants is not unexpected because this is the most common foraging guild among birds worldwide. Overall, we found no prevalence of any specific foraging guild among altitudinal migrants across zoogeographic regions. The results of studies to date suggest that altitudinal migration by birds may be driven by a number of factors, including access to increased food resources for breeding or molting, weather conditions, and mating and nesting opportunities. However, to better understand the mechanisms underlying altitudinal migration, broadening the geographic scope of studies is paramount, with additional study of altitudinal migration especially needed in the megadiverse tropical regions of sub‐Saharan Africa, Southeast Asia, and South America.     

Diffuse, short and slow migration among Blue Tits

The knowledge of migration systems in long-distance regular migrants is in many cases extensive. Our understanding of the migratory characteristics of partial migrants, on the other hand, is far more rudimentary. We investigated migratory characteristics of partially migratory Blue Tits Cyanistes caeruleus using ringing recoveries of Swedish birds, to answer questions about geographic migration patterns, age-specific migrations, migration speeds and synchrony of movements. Median migration distance of Swedish Blue Tits was 82 km, with a main autumn direction in the sector between S and W (large directional scatter). Northerly and southerly populations did not differ in migration directions or distances, suggesting chain migration to be the general pattern. A larger proportion of adult Blue Tits remained near the breeding grounds during winter than was the case for juveniles. Some of the migrating birds (17%) seemed not to return in spring but stayed to breed closer to the winter area. Swedish Blue Tits show an exceptionally slow migration speed (median 13 km/day), among the slowest speeds recorded for any migrant bird. The Blue Tit represents an extreme case of diffuse, short and slow bird migration.     

Effects of migration distance and sex on stopover timing and refueling by Wilson's Warblers

Migration distance and sex are integral to avian migration strategies, yet these intrinsic factors are understudied with respect to their effects on stopover ecology and behavior. We investigated how individual variation in migration distance and sex affected body condition, refueling performance, and stopover durations of Wilson's Warblers (Cardellina pusilla) during spring and fall 2006 at a stopover site in southern Arizona. Migration distance of individuals was inferred using the hydrogen stable isotope ratio of feathers (δ²H_f) as an index of breeding latitude, refueling rate was assessed using plasma triglyceride and β‐OH‐butyrate, and minimum length of stay was estimated by re‐sighting color‐banded birds. In the spring, migration distance and sex were strong determinants of the timing of migration by Wilson's Warblers, with males and shorter‐distance migrants passing through the site earlier than females and longer‐distance migrants. Later‐arriving migrants also had higher fat scores and refueling rates. However, neither migration distance nor sex independently affected body condition (fat score or size‐corrected mass), refueling rate, or minimum length of stay. In a smaller sample of fall migrants, we found that longer‐distance migrants and males had higher refueling rates than shorter‐distance migrants and females. Our results show that differences in migration distance can, under some circumstances, affect how birds use stopover sites. Stable isotope analysis and other methods can provide information on migration distance, and should be integrated with measurements of timing, fuel stores, refueling performance, and departure behavior to gain a deeper understanding of bird migration.     

Seasonal differences in behavior patterns of the migratory white-throated sparrow

Avian migration and associated changes in behavior patterns occur on a seasonal basis and are regulated by both circadian and circannual clocks. It is well known that seasonally migrating birds must alter the temporal partitioning of behaviors in order to successfully complete nocturnal migration, although the precise changes in allocation have not previously been examined. Because of the difficulties related to the study of bird migration in the wild, many studies regarding the physiology of migration have been performed in captivity, where nocturnal migratory restlessness is readily observable at the appropriate times of the year. Even so, quantifying the amount and timing of daily behavioral rhythms in migrating and non-migrating individuals has not previously been performed. We used video recordings of a common nocturnal migrant, the white-throated sparrow (Zonotrichia albicollis), to generate a basic ethogram and to chart the timing and frequency of occurrence of each behavior in order to compare migrant individuals with non-migrating birds. Our study suggests that the specific allocation of behaviors are altered during migration and that these changes might contribute to the development of the “migratory syndrome”.