Layers of nocturnal insect migrants at high-altitude: the influence of atmospheric conditions on their formation

• 1 Radar studies of nocturnal insect migration have often found that the migrants tend to form well‐defined horizontal layers at a particular altitude.
• 2 In previous short‐term studies, nocturnal layers were usually observed to occur at the same altitude as certain meteorological features, most notably at the altitudes of temperature inversion tops or nocturnal wind jets.
• 3 Statistical analyses are presented of 4 years of data that compared the presence, sharpness and duration of nocturnal layer profiles, observed using continuously‐operating entomological radar, with meteorological variables at typical layer altitudes over the U.K.
• 4 Analysis of these large datasets demonstrated that temperature was the foremost meteorological factor that was persistently associated with the presence and formation of longer‐lasting and sharper layers of migrating insects over southern U.K.

     

Long‐range seasonal migration in insects: mechanisms,evolutionary drivers and ecological consequences

Myriad tiny insect species take to the air to engage in windborne migration, but entomology also has its ‘charismatic megafauna’ of butterflies, large moths, dragonflies and locusts. The spectacular migrations of large day‐flying insects have long fascinated humankind, and since the advent of radar entomology much has been revealed about high‐altitude night‐time insect migrations. Over the last decade, there have been significant advances in insect migration research, which we review here. In particular, we highlight: (1) notable improvements in our understanding of lepidopteran navigation strategies, including the hitherto unsuspected capabilities of high‐altitude migrants to select favourable winds and orientate adaptively, (2) progress in unravelling the neuronal mechanisms underlying sun compass orientation and in identifying the genetic complex underpinning key traits associated with migration behaviour and performance in the monarch butterfly, and (3) improvements in our knowledge of the multifaceted interactions between disease agents and insect migrants, in terms of direct effects on migration success and pathogen spread, and indirect effects on the evolution of migratory systems. We conclude by highlighting the progress that can be made through inter‐phyla comparisons, and identify future research areas that will enhance our understanding of insect migration strategies within an eco‐evolutionary perspective.     

Flight altitude of trans-Sahara migrants in autumn: a comparison of radar observations with predictions from meteorological conditions and water and energy balance models

Radar observations on the altitude of bird migration and altitudinal profiles of meteorological conditions over the Sahara desert are presented for the autumn migratory period. Migratory birds fly at an average altitude of 1016 m (a.s.l.) during the day and 571 m during the night. Weather data served to calculate flight range using two models: an energy model (EM) and an energy-and-water model (EWM). The EM assumes that fuel supply limits flight range whereas the EWM assumes that both fuel and water may limit flight range. Flight ranges estimated with the EM were generally longer than those with the EWM. This indicates that trans-Sahara migrants might have more problems balancing their water than their energy budget. However, if we assume fuel stores to consist of 70% instead of 100% fat (the remainder consisting of 9% protein and 21% water), predicted flight ranges of the EM and EWM largely overlap. Increased oxygen extraction, reduced flight costs, reduced exhaled air temperature, reduced cutaneous water loss and increased tolerance to water loss are potential physiological adaptations that would improve the water budget in migrants. Both the EM and EWM predict optimal flight altitudes in agreement with radar observations in autumn. Optimal flight altitudes are differently predicted by the EM and EWM for nocturnal spring migration. During spring, the EWM predicts moderately higher and the EM substantially higher flight altitudes than during autumn. EWM predictions are therefore in better agreement with radar observations on flight altitude of migrants over the Negev desert in spring than EM predictions.     

High Altitude Bird Migration at Temperate Latitudes: A Synoptic Perspective on Wind Assistance

At temperate latitudes the synoptic patterns of bird migration are strongly structured by the presence of cyclones and anticyclones, both in the horizontal and altitudinal dimensions. In certain synoptic conditions, birds may efficiently cross regions with opposing surface wind by choosing a higher flight altitude with more favourable wind. We observed migratory passerines at mid-latitudes that selected high altitude wind optima on particular nights, leading to the formation of structured migration layers at varying altitude up to 3 km. Using long-term vertical profiling of bird migration by C-band Doppler radar in the Netherlands, we find that such migration layers occur nearly exclusively during spring migration in the presence of a high-pressure system. A conceptual analytic framework providing insight into the synoptic patterns of wind assistance for migrants that includes the altitudinal dimension has so far been lacking. We present a simple model for a baroclinic atmosphere that relates vertical profiles of wind assistance to the pressure and temperature patterns occurring at temperate latitudes. We show how the magnitude and direction of the large scale horizontal temperature gradient affects the relative gain in wind assistance that migrants obtain through ascending. Temperature gradients typical for northerly high-pressure systems in spring are shown to cause high altitude wind optima in the easterly sectors of anticyclones, thereby explaining the frequent observations of high altitude migration in these synoptic conditions. Given the recurring synoptic arrangements of pressure systems across temperate continents, the opportunities for exploiting high altitude wind will differ between flyways, for example between easterly and westerly oceanic coasts.     

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.     

东北地区粘虫的季节性迁飞行为(英文)

粘虫Mythimna separata是我国农业生产上的重要害虫, 为了明确其季节性迁飞行为参数, 本研究采用垂直监测昆虫雷达（vertical-looking radar, VLR）及相关辅助设备的长期自动观测, 结合基于GIS的大区环流和轨迹模拟, 调查分析了2005年东北地区粘虫季节性迁飞行为。结果表明： 粘虫在不同季节和夜间不同时间空中飞行高度具有明显差异, 空中飞行行为受气象条件尤其是空中风场影响较大; 春季和秋季主要借助气流运载进行大规模长距离迁飞, 夜间持续飞行时间可达9 h, 多数个体能完成整夜飞行, 春季迁飞高度主要在300~600 m, 秋季飞行高度相对较低主要在300 m以下和400~500 m。夏季雷达回波有明显的成层现象, 最高可达1 000 m, 主要集中在500 m和700 m两个高度层。轨迹分析显示： 5月29日由山东潍坊、 临沂等虫源地起飞的黏虫, 顺西南气流越海迁飞, 6月1日在气旋天气影响下, 在吉林省白城等地降落; 7月中旬主要为当地黏虫受对流天气影响进行短距离迁飞扩散; 9月11日虫源来自内蒙古呼伦贝尔, 顺西北气流向吉林省东南方向迁飞。研究结果为东北地区粘虫的有效防控提供了技术支撑。     

The long-distance migration of Nilaparvata lugens (Stål) (Delphacidae) in China: radar observations of mass return flight in the autumn

Abstract.

• 1 A case study is presented of the autumn migration of the brown planthopper, Nilaparvata lugens (Stål), in the area of Nanjing in the People's Republic of China. The study was made using a high frequency (8 mm wavelength) radar and a net suspended from a kytoon.
• 2 The observations confirmed that long-distance return migrations occur in China in mid and late September, with N.lugens being carried on the prevailing north-easterly wind towards the autumn infestation and overwintering areas of the species.
• 3 After mass take-off in the late afternoon or at dusk, the migrants flew for several hours during the evening, often in a dense layer which formed at heights between about 400 and 1000m above ground. These layers often had well-defined ceilings corresponding to an air temperature of about 16°C. The migration height was above the top of the surface temperature inversion, i.e. the migrants did not fly at the height of the warmest air.
• 4 The dense layer concentrations overflying the radar were backtracked to source areas up to 240 km away in the north-east of Jiangsu Province. Planthoppers observed emigrating from the Nanjing area would reach areas in south Anhui Province or north Jiangxi Province if they flew for 12 h.
• 5 There was a second period of mass take-off at dawn. Insect layers sometimes formed but did not last longer than 1–2h.
• 6 The present results were strikingly different from those previously observed in the dry season in the Philippines, where migratory flight durations were largely confined to periods of about 30min at dusk and dawn.
• 7 Our observations are discussed in relation to the equator-wards return migrations undertaken in autumn by other insect species, and the importance of these migrations for the maintenance of long-flying genotypes in the overwintering populations is considered.

     

NUMERICAL SIMULATION OF THE PATHWAYS OF MIGRATING INSECTS

Abstract  Based on boundary layer meteorology and behavioural ecology of insect migration, a numerical TRA model was established. The spatial and temporal resolutions of the standard meteorological data are far insufficient for insect migration studies; so, a one dimension TKE closure scheme (E-ε scheme) was adopted to simulate the wind and temperature profiles en route based on the conventional surface and 850 hPa wind and temperature data of Chinese National Meteorological Bureau. Then the migrating behaviour of insects was parameterized as some proper mathematical expressions to determine their timing and their flight height, speed and direction from the wind and temperature profiles simulated by the PBL sub-model, and so their flight pathways could be estimated by a simple algorithm. Finally, the hourly episodes of the airborne migrants were output which consists of the latitude, longitude and flying altitude. The TRA model was verified by mark-release-recapture studies of Mythimna separata, Loxostege sticticalis , and Agrotis ipsilon . The results suggest that the parameterizing scheme of migratory behaviour, the numerical simulation scheme of wind and temperature in PBL and the TRA procedure developed in this paper should be reasonable and feasible. This model provides a useful tool for inter-regional forecast of migratory insect pests.     

Radar observations of moths migrating in a nocturnal low-level jet

Abstract. 1. Radar observations of insects migrating at night over central-western New South Wales have detected an instance of migration in a low-level wind jet.
2. From the characteristics of the radar echoes, and from the catches obtained in traps at ground level and at the altitude of migration, the migrants can be identified as noctuid and pyralid moths of a number of different species.
3. The migration, which was in a downwind direction, started at dusk and ended at about dawn. During the period immediately before first light, a large proportion of the migrants were concentrated into a 100m deep layer at an altitude of about 250m; this layer had not been present during the first half of the night.
4. The boundary layer wind profile at dawn exhibited a clear low-level jet structure, with a wind maximum between 100 and 300m, and strong shear in the wind direction below 300m. A strong surface temperature inversion, but not a wind-speed maximum, had been present the previous evening.
5. The formation of the layer concentration in the upper part of the jet may be accounted for in terms of previously described responses of nocturnally migrating insects to a surface temperature inversion. It is not therefore necessary to assume that the migrants were responding specifically to the presence of a wind-speed maximum.     

Monitoring bird migration with a fixed-beam radar and a thermal-imaging camera

ABSTRACT.   Previous studies using thermal imaging cameras (TI) have used target size as an indicator of target altitude when radar was not available, but this approach may lead to errors if birds that differ greatly in size are actually flying at the same altitude. To overcome this potential difficulty and obtain more accurate measures of the flight altitudes and numbers of individual migrants, we have developed a technique that combines a vertically pointed stationary radar beam and a vertically pointed thermal imaging camera (VERTRAD/TI). The TI provides accurate counts of the birds passing through a fixed, circular sampling area in the TI display, and the radar provides accurate data on their flight altitudes. We analyzed samples of VERTRAD/TI video data collected during nocturnal fall migration in 2000 and 2003 and during the arrival of spring trans-Gulf migration during the daytime in 2003. We used a video peak store (VPS) to make time exposures of target tracks in the video record of the TI and developed criteria to distinguish birds, foraging bats, and insects based on characteristics of the tracks in the VPS images and the altitude of the targets. The TI worked equally well during daytime and nighttime observations and best when skies were clear, because thermal radiance from cloud heat often obscured targets. The VERTRAD/TI system, though costly, is a valuable tool for measuring accurate bird migration traffic rates (the number of birds crossing 1609.34 m [1 statute mile] of front per hour) for different altitudinal strata above 25 m. The technique can be used to estimate the potential risk of migrating birds colliding with man-made obstacles of various heights (e.g., communication and broadcast towers and wind turbines)—a subject of increasing importance to conservation biologists.     

Doppler radar detection of exceptional mass-migration of aphids into Finland

Our objective was to detect mass migrations of insects of economic significance by insect traps and a Doppler weather radar. Migrants were sampled by suction traps, tow nets and light traps in the Helsinki region. We used radar to observe the migrating insects, and trajectories to backtrack mass migrations of aphids (Homoptera, Aphididae) in spring 1988. The aphid migrations were clearly observed in trap catches and by radar. The first migration, mainly involving Euceraphis betulae, occurred on 18 May and was tracked back to northern Poland. The second migration, mainly of Rhopalosiphum padi (a serious pest of small-grain cereals), occurred 3 days later and was tracked back to a large area covering Latvia and western Russia south of St Petersburg. The third migration included both E. betulae and R. padi, and took place on 30 May. It originated from Estonia. Neither trap nor radar data provide exact quantitative information on migrations. Trapping efficiency depends strongly on wind speed and insect size. Radar echo intensity is very strongly related to the sizes of insects in the large volume of air measured, and the sizes are not known accurately. Weather data, especially temperature, can be used in predicting the development of aphids, and air-parcel trajectories in estimating the source areas of migrants. These methods for forecasting aphid migrations, combined with radar observations, are useful for warning purposes and to intensify insect trapping. This would contribute to more efficient agricultural pest management. Received: 11 March 2000 / Revised: 24 April 2000 / Accepted: 26 April 2000     

Spring passerine migrants stopping over in the Sahara are not fall-outs

The strategy of migrants crossing the Sahara desert has been the subject of debate, but recent evidence from radar studies has confirmed that most passerines use an intermittent migration strategy. The latter has also been suggested from previous studies in oases during autumn migration. It was found that migrants with relatively high fuel loads rest in the desert during daytime and continue migration during the following night, whereas lean migrants stopover in oases for several days to refuel. However, data from the Sahara are scarce for spring migration. We captured passerine migrants near B?r Amrane (22°47′N, 8°43′W) in the plain desert of Mauritania for 3 weeks during spring migration in 2004. We estimated flight ranges of 85 passerines stopping over in the desert to test whether they carried sufficient fuel loads to accomplish migration across the Sahara successfully. High fat loads of the majority of birds indicated that they were neither “fall-outs” nor too weak to accomplish migration successfully. The flight range estimates, based on mean flight speeds derived from radar measurements (59 km/h), revealed that 85% of all birds were able to reach the northern fringe of the desert with an intermittent migration strategy. Furthermore, birds stopping over in an oasis (Ouadane, 370 km to the southwest of B?r Amrane) did not carry consistently lower fuel loads compared to the migrants captured in the desert.     

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.     

The flight behaviour of individual passerine nocturnal migrants: A tracking radar study

Nocturnal passerine migrants were tracked with a small automatic tracking radar during spring migration in eastern New York. Climbing, descending and markedly non-linear tracks were selected for analysis because they may reveal relationships not evident in normal straight and level tracks. Climbing individuals ascended at 1 to 2 vertical metres per second by heading into the wind and increasing their ascent angles while air speed tended to remain constant. Within individual tracks, birds flew slower when flying downwind than when flying into the wind and changes in air speed were performed over periods of a few seconds. A small amount of data suggested that this behaviour did not occur under overcast skies. Both the direction and speed of the wind force were important in predicting air speed. Multiple regression analysis indicated that faster flying birds were more likely to fly in winds of high speed and at large angles into the wind.     

Altitude choice by night migrants in a desert area predicted by meteorological factors

The height distribution of nocturnal migrants in southern Israel was determined by con-ically scanning the sky with the pencil-beam of an X-band radar at different elevation angles. Altitudinal profiles of meteorological parameters were derived from radio sondes launched at midnight and from pilot balloons launched every 4 h. A model to predict the height distribution of birds by means of meteorological variables was developed by assuming that the observed proportions of birds within a height zone, compared with the neighbouring height zones, reflect the degree of the birds' preference for that height zone. Only one among the variables included in the multiple regression analysis proved to have a significant influence on the height distribution of migrants: the difference of tailwind speed between height zones. Simulations with 1000 birds choosing altitudes by means of the night's altitudinal profile of tailwind speed closely traced the observed distributions. The fact that all the other meteorological factors which were previously suggested to have an influence on the flight range in trans-desert migration were not selected as relevant factors is discussed. The following basic information on nocturnal bird migration in the Negev is provided as a background for the statistical analysis: Directions of migration are within very narrow limits. During the first hour after take-off, 60% of autumn migrants and 75% of spring migrants are climbing, with vertical speeds of 0.1–2 m per s and 0.1–4 m per s, respectively. During the rest of the night, climbing and descending birds are in nearly equal proportions. Thus, there is a high potential of sampling atmospheric conditions at different altitudes. Height distributions in spring and autumn show the influence of the trade wind situation, autumn migrants making use of the northerly winds at low levels in spite of high temperatures, while spring migrants tend to reach the southwesterly winds at higher levels.     

Multi‐generational long‐distance migration of insects: studying the painted lady butterfly in the Western Palaearctic

Long‐range, seasonal migration is a widespread phenomenon among insects, allowing them to track and exploit abundant but ephemeral resources over vast geographical areas. However, the basic patterns of how species shift across multiple locations and seasons are unknown in most cases, even though migrant species comprise an important component of the temperate‐zone biota. The painted lady butterfly Vanessa cardui is such an example; a cosmopolitan continuously‐brooded species which migrates each year between Africa and Europe, sometimes in enormous numbers. The migration of 2009 was one of the most impressive recorded, and thousands of observations were collected through citizen science programmes and systematic entomological surveys, such as high altitude insect‐monitoring radar and ground‐based butterfly monitoring schemes. Here we use V. cardui as a model species to better understand insect migration in the Western Palaearctic, and we capitalise on the complementary data sources available for this iconic butterfly. The migratory cycle in this species involves six generations, encompassing a latitudinal shift of thousands of kilometres (up to 60 degrees of latitude). The cycle comprises an annual poleward advance of the populations in spring followed by an equatorward return movement in autumn, with returning individuals potentially flying thousands of kilometres. We show that many long‐distance migrants take advantage of favourable winds, moving downwind at high elevation (from some tens of metres from the ground to altitudes over 1000 m), pointing at strong similarities in the flight strategies used by V. cardui and other migrant Lepidoptera. Our results reveal the highly successful strategy that has evolved in these insects, and provide a useful framework for a better understanding of long‐distance seasonal migration in the temperate regions worldwide.     

昆虫迁飞行为的参数化Ⅰ.行为分析

通过对雷达昆虫学研究和其他方法得到的研究成果的综合分析,提出一套昆虫迁飞行为参数化方案。即：起飞时间以日出日没及晨昏朦影时刻为基准,降落时间依目标昆虫的迁飞特性具体取值;运行高度取边界层顶与目标昆虫飞行低温阈限所在高度之间的气流层,运行方向取风向值并以目标昆虫的定向加以修饰,运行速度为风速与目标昆虫自身飞行速度的矢量和。这套方案可作为昆虫迁飞轨迹数值模拟的基础,为迁飞性害虫异地预测提供一种有效的工具     

QUANTITATIVE STUDY OF MIGRATION WITH 23-CENTIMETRE RADAR*

A quantitative study was made of the displays of "angels" on the screen of a 23-centimetre radar installation on Cape Cod, Massachusetts. The birds responsible for the displays were nocturnal migrants, mainly passerines.
Other things being equal, the density of angels decreased roughly exponentially with distance from the centre of the radar screen, being halved about every eight miles. The rate of decrease varied only slightly with total migration density, and could be used to estimate the total number of bird targets in the air.
Estimates obtained by this means were compared with estimates of migration density obtained from "moon-watching" observations, which were reduced by a new method of calculation which incorporates radar measurements of the height of flight. Each angel was found to correspond to between two and twelve birds. After considering and rejecting other explanations, it is concluded that many birds migrate in groups at night. There is evidence that many species (including Parulidae, Turdidae and Emberizidae) migrate in small groups, rather than a few species in large groups. Groups of passerine night-migrants are probably spread over a wide area, since it is rare to see two birds cross the moon together.
Once the average group-size has been determined in this way, radar can be used to measure the density of migration over a large area. Estimates of low migration density have a standard error about 25%, but estimates of high migration density are less accurate, and very high migration densities cannot be measured at all, although they can be accurately identified as such.
The results in this paper cannot be applied directly to other radar stations, nor to other species of birds.     

Convergent patterns of long-distance nocturnal migration in noctuid moths and passerine birds

Vast numbers of insects and passerines achieve long-distance migrations between summer and winter locations by undertaking high-altitude nocturnal flights. Insects such as noctuid moths fly relatively slowly in relation to the surrounding air, with airspeeds approximately one-third of that of passerines. Thus, it has been widely assumed that windborne insect migrants will have comparatively little control over their migration speed and direction compared with migrant birds. We used radar to carry out the first comparative analyses of the flight behaviour and migratory strategies of insects and birds under nearly equivalent natural conditions. Contrary to expectations, noctuid moths attained almost identical ground speeds and travel directions compared with passerines, despite their very different flight powers and sensory capacities. Moths achieved fast travel speeds in seasonally appropriate migration directions by exploiting favourably directed winds and selecting flight altitudes that coincided with the fastest air streams. By contrast, passerines were less selective of wind conditions, relying on self-powered flight in their seasonally preferred direction, often with little or no tailwind assistance. Our results demonstrate that noctuid moths and passerines show contrasting risk-prone and risk-averse migratory strategies in relation to wind. Comparative studies of the flight behaviours of distantly related taxa are critically important for understanding the evolution of animal migration strategies.     

10. PROCEEDINGS OF THE SEVENTEENTH ANNUAL GENERAL MEETING OF THE S.A. ORNITHOLOGICAL SOCIETY HELD IN THE COMMITTEE ROOM OF THE PUBLIC LIBRARY. JOHANNESBURG,AT 3 P.M., SATURDAY, 8TH MARCH. 1947

Bruderer, B. 1994. Nocturnal bud migration in the Negev (Israel) a tracking radar study. Ostrich 65: 204–212.

The present publication summarizes the methodological possibilities of tracking radar and describes some features of nocturnal migration at two sites in the Negev, which include anwers to basic questions of bird migration. The directions of spring and autumn migration were practically opposite; only the headings in spring indicated some more compensation for stronger westerly winds. The volume of nocturnal spring migration was only about 65% of autumn migration, which may be an indication of mortality outside tie breeding area. Highest densities of migration at the two radar sites in the Negev Highlands (450 m above sea level) and in the Arava Valley (150 m below sea level) indicated flightlevels adjusted to atmospheric conditions aloft, and not to round level. Due to the trade-wind system, the birds heading southward in autumn flew mainly below flew mainly above 1500 m above sea level, while in spring they tended to make use of the anti-trades at higher altitudes. The decisive factor for altitude choice was the speed of tailwind in spring and autumn; other factors, such as temperature, humidity and pressure had no significant influence on the altitude distributions. With respect to the question of non-stop or intermittent flight across large desert areas, the data show that between the eastern deserts of Egypt and the Sinai/Negev complex the nocturnal migrants maintained their schedule of nocturnal flight and diurnal rest. A few exceptions of nocturnal migrants continuing migration at high altitudes into the day were identified mainly as heron- and gull-type birds. The proportion of waders and waterfowl identified by wing-beat pattern in nocturnal migration is nearly the same at both sites, indicating broad-front migration across the desert. The numbers of birds with continuous wingbeats is, however, so large compared to available estimates of waders and waterfowl wintering in Africa that careful reconsideration of the underlying assumptions in the radar and field estimates is necessary.