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.     

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.     

Traveling or stopping of migrating birds in relation to wind: an illustration for the osprey

Although it is often assumed that birds strongly prefer tailwindsfor their migratory flights, we predict that a strategy of nowind selectivity (traveling independently of winds) may be morefavorable than wind selectivity (traveling on tailwind occasionsbut stopping to rest under headwind occasions) for birds withlow energy costs of travel relative to rest and for birds thatcannot use stopover time for efficient fuel deposition. We testthis prediction by analyzing the daily traveling or stoppingas recorded by satellite tracking of five ospreys Pandion haliaetus,a species often using energy-saving thermal soaring, duringtheir migration between northern Europe and Africa. Besideswind, precipitation is another weather factor included in theanalyses because thermal soaring migrants are expected to stopand rest in rainy weather. In logistic regression analyses,taking into account the effects of latitude, behavior on previousday, season, date, and individual for discriminating betweentraveling and stopping days, we found a lack of influence ofwinds, suggesting that the ospreys travel or stop without regardto wind. This lack of wind selectivity under light and moderatewinds is in agreement with our prediction. We expect a low degreeof wind selectivity and thus regular flights under headwindsalso among other types of birds that cannot use stopping timefor efficient foraging and fuel deposition. We also found anunexpected lack of influence of precipitation, possibly becauseof relatively few instances with rainfall in combination withpoor geographic precision for estimates of this weather variable.     

Can wind help explain seasonal differences in avian migration speed?

A bird's ground speed is influenced by the wind conditions it encounters. Wind conditions, although variable, are not entirely random. Instead, wind exhibits persistent spatial and temporal dynamics described by the general circulation of the atmosphere. As such, in certain geographical areas wind's assistance (or hindrance) on migratory flight is also persistent, being dependent upon the bird's migratory direction in relation to prevailing wind conditions. We propose that, considering the western migration route of nocturnal migrants through Europe, winds should be more supportive in spring than in autumn. Thus, we expect higher ground speeds, contributing to higher overall migration speeds, in spring. To test whether winds were more supportive in spring than autumn, we quantified monthly wind conditions within western Europe relative to the seasonal direction of migration using 30 years (1978–2008) of wind data from the NCEP/NCAR Reanalysis dataset. We found that supporting winds were significantly more frequent for spring migration compared to autumn and up to twice as frequent at higher altitudes. We then analyzed three years (2006–2008) of nocturnal migratory ground speeds measured with radar in the Netherlands which confirmed higher ground speeds in spring than autumn. This seasonal difference in ground speed suggests a 16.9% increase in migration speed from autumn to spring. These results stress the importance of considering the specific wind conditions experienced by birds when interpreting migration speed. We provide a simple methodological approach enabling researchers to quantify regional wind conditions for any geographic area and time period of interest.     

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.     

Morning flight behavior of nocturnally migrating birds along the western basin of Lake Erie

Many species of birds that normally migrate during the night have been observed engaging in so‐called morning flights during the early morning. The results of previous studies have supported the hypothesis that one function of morning flights is to compensate for wind drift that birds experienced during the night. Our objective was to further explore this hypothesis in a unique geographic context. We determined the orientation of morning flights along the southern shore of Lake Erie's western basin during the spring migrations of 2016 and 2017. This orientation was then compared to the observed orientation of nocturnal migration. Additionally, the orientation of the birds engaged in morning flights following nights with drifting winds was compared to that of birds following nights with non‐drifting winds. The morning flights of most birds at our observation site were oriented to the west‐northwest, following the southern coast of Lake Erie. Given that nocturnal migration was oriented generally east of north, the orientation of morning flight necessarily reflected compensation for accumulated, seasonal wind drift resulting from prevailingly westerly winds. However, the orientation of morning flights was similar following nights with drifting and non‐drifting winds, suggesting that birds on any given morning were not necessarily re‐orienting as an immediate response to drift that occurred the previous night. Given the topographical characteristics of our observation area, the west‐northwest movement of birds in our study is likely best explained as a more complex interaction that could include some combination of compensation for wind drift, a search for suitable stopover habitat, flying in a direction that minimizes any loss in progressing northward toward the migratory goal, and avoidance of a lake crossing.     

Great flights by great snipes: long and fast non-stop migration over benign habitats

Migratory land birds perform extreme endurance flights when crossing ecological barriers, such as deserts, oceans and ice-caps. When travelling over benign areas, birds are expected to migrate by shorter flight steps, since carrying the heavy fuel loads needed for long non-stop flights comes at considerable cost. Here, we show that great snipes Gallinago media made long and fast non-stop flights (4300-6800 km in 48-96 h), not only over deserts and seas but also over wide areas of suitable habitats, which represents a previously unknown migration strategy among land birds. Furthermore, the great snipes achieved very high ground speeds (15-27 m s(-1)), which was not an effect of strong tailwind support, and we know of no other animal that travels this rapidly over such a long distance. Our results demonstrate that some migratory birds are prepared to accept extreme costs of strenuous exercise and large fuel loads, even when stopover sites are available along the route and there is little tailwind assistance. A strategy of storing a lot of energy before departure, even if migration is over benign habitats, may be advantageous owing to differential conditions of fuel deposition, predation or infection risk along the migration route.     

Nocturnal migratory flight initiation in reed warblers Acrocephalus scirpaceus: effect of wind on orientation and timing of migration

We used radio-telemetry to study autumn migratory flight initiation and orientation in relation to wind and air pressure in a nocturnal passerine migrant, the reed warbler Acrocephalus scirpaceus at Falsterbo, southwest Sweden. The majority of the reed warblers departed in the expected migratory direction towards south of southwest, while a low number of the birds took off in reverse directions between north and east. Flight directions at departure correlated with wind directions. These correlations were particularly prominent at higher wind speeds but were absent at wind speeds below 4 m/s. Birds departing in the expected migratory direction compensated completely for wind drift. The reed warblers preferred to depart during nights with tailwinds and when air pressure was increasing suggesting that reed warblers are sensitive to winds and air pressure and select favourable wind conditions for their migratory flights. Since air pressure as well as velocity and direction of the wind are correlated with the passage of cyclones, a combination of these weather variables is presumably important for the birds' decision to migrate and should therefore be considered in optimal migration models.     

Field studies of avian nocturnal migratory orientation I. interaction of sun,wind and stars as directional cues

Tracking radar and visual observation techniques were used to observe the orientation of free-flying passerine nocturnal migrants in situations in which potentially usable directional cues were absent or gave conflicting information. When migrants had seen the sun near the time of sunset and/or the stars, they oriented in appropriate migratory directions even when winds were opposed. Under solid overcast skies that prevented a view of both sun and stars, the birds headed downwind in opposing winds and thus moved in seasonally inappropriate directions. The data point to the primacy of visual cues over wind direction, with either sun or stars being sufficient to allow the birds to determine the appropriate migration direction.     

Environmental influences on the orientation of free-flying nocturnal bird migrants

Autumn nocturnal bird migration was investigated using radar and visual observations. Multivariate analyses assessed the influence of environmental variables on orientation. Two categories of birds were distinguished: (1) shorebirds and waterfowl migrating in flocks; and (2) passerine birds flying singly. These two classes of migrants employ different orientation mechanisms. Landbirds selectively flew with the wind, regardless of its direction or speed or whether the skies were clear or overcast. Shorebirds and waterfowl flew in directions independent of wind in light or moderate winds. The accuracy of passerine orientation was not correlated with any of the variables examined. The passerine orientation observed cannot be accounted for by stellar orientation, but is explicable via orientation on the basis of wind direction.     

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.     

A RADAR STUDY OF THE AUTUMN MIGRATION OF WOOD PIGEONS COLUMBA PALUMBUS IN SOUTHERN SCANDINAVIA

The migration of Wood Pigeons in southern Scandinavia was studied from 21 September to 10 October 1971 and from 16 September to 15 November 1972 using radar stations supplemented with observations from an aircraft and a network of ground observers. By far the largest quantities of Wood Pigeons migrated after cold front passages with northwesterly to northeasterly tailwinds. Most birds departed on a few days, apparently as a consequence of strong preference for tailwind situations. With northwesterly winds a proportionately high migratory activity was recorded in the Kattegatt area. With northeasterly winds activity was higher in the Baltic area. This allowed the Wood Pigeons to make maximal use of the tailwind component, and their ground speed usually exceeded 80 km/h. The calculated mean air speed was 60 km/h. Their dependence on tailwind was particularly strong when the birds were engaging in long sea-crossings, such as across the Kattegatt. Different coastlines affected the geographical pattern of migration in different ways. Frequently Wood Pigeon flocks flew almost parallel to the coast but some distance off shore, until they finally departed. The deflective force of coastlines was greatest when the birds' ground speed was low, that is, under headwind conditions or in calm weather. Mean track directions measured over two areas in northern Skane, called Inland W and Inland E, situated about 60 km apart, differed by 11, those over the western area being directed more to the south than those over the eastern. No significant correlation with wind directions was found in these areas. Combining data from the whole land area, however, track directions were found to vary from day to day in significant correlation to the wind direction. Mean track directions over the Baltic agreed with those over Skane, but both differed significantly from those over the Kattegatt. Both over the Baltic and over the Kattegatt directions were significantly correlated with wind directions, and showed greater variation than track directions over land. Daily track differences over the Baltic resulted both from differences taking place over the land, and from real wind deflection (drift). Both over the land and over the sea heading directions were correlated with wind directions, suggesting compensatory efforts on the part of the birds. On three days extensive fog covered much of the study area. Wood Pigeons continued to migrate, but certain aberrations in their behaviour were noted. Over land migration was relatively heavier in the west with northwesterly winds and in the east with northeasterly winds. The correlation demonstrated between wind direction and the mean track direction was based upon the fact that populations with different inherent primary directions made up different proportions of the migrating cohorts under different wind conditions (pseudo-drift). The incomplete compensation for wind deflection over the sea is ascribed to the lack of visual orientation cues. The more accurate orientation possible over land suggests one reason for the birds' reluctance to flights across the open sea. When mean track directions of Wood Pigeons in different parts of southern Scandinavia were related to the migratory goals of these birds, it was found that they have to change their primary direction in the course of their journey from breeding to wintering areas.     

Migration and orientation of passerine night migrants in northeast England

Evening departures of passerine migrants were watched by radar in northeast England on 193 nights in the months of August to November 1961 to 1963. It was found that migrants were not departing each night on the same heading, from which they were drifted passively by the wind, but rather were departing on similar tracks night after night. On ten nights they maintained their tracks even when the upper-air wind changed during the night. This indicates that migrants can, at least sometimes, compensate completely for the effect of the wind and fly on a fixed track, but calculations show that there are certain wind conditions when this becomes impossible.
Movements were grouped according to the months in which they took place and the tracks which they followed, and then related to variousweather factors. Migration was seen most frequently with light winds favourable to the birds'track, after a marked drop in temperature (in October/November, but not August/September), and withabsence of cloud. However, it was also recorded on 17 nights when the cloud cover was complete.
The departures of small passerines from Britain in August and September follow tracks east of south, yet the same birds reach western Iberia later in their journey. It is suggested that the direction of departure from Britain has been evolved to take advantage of the prevailing (westerly) upper-air winds. Certain warblers leave their breeding areas in Northumberland without laying down much migratory fat, yet the same species make long flights non-stop (across the Mediterranean and Sahara) later in their migrations. These migrants change froman insect to a fruit diet in autumn, and can presumably IaKdown fat more quickly the further south they travel in Europe, as the soft fruit ripens earlier in southern areas.     

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.     

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 migratory songbirds adjust their travelling direction aloft: evidence from a radiotelemetry and radar study

In order to fully understand the orientation behaviour of migrating birds, it is important to understand when birds set their travel direction. Departure directions of migratory passerines leaving stopover sites are often assumed to reflect the birds'' intended travel directions, but this assumption has not been critically tested. We used data from an automated radiotelemetry system and a tracking radar at Falsterbo peninsula, Sweden, to compare the initial orientation of departing songbirds (recorded by radiotelemetry) with the orientation of songbird migrants in climbing and level flight (recorded by radar). We found that the track directions of birds at high altitudes and in level flight were more concentrated than the directions of departing birds and birds in climbing flight, which indicates that the birds adjust their travelling direction once aloft. This was further supported by a wide scatter of vanishing bearings in a subsample of radio-tracked birds that later passed an offshore radio receiver station 50 km southeast of Falsterbo. Track directions seemed to be more affected by winds in climbing compared with level flights, which may be explained by birds not starting to partially compensate for wind drift until they have reached cruising altitudes.     

Stopover and fat deposition by North American wood-warblers (Parulinae) following spring migration over the Gulf of Mexico

Summary Length of stopover and rate of weight gain (fat deposition) were studied in several species of passerine birds that stopped in southwestern Louisiana along the northern coast of the Gulf of Mexico after a trans-Gulf flight. Fatdepleted birds were more common among the birds that arrived at our study site in southwest Louisiana, though variability characterized our samples. Migrants that landed after encountering opposing winds or rain over the northern Gulf of Mexico were, on average, fatter than migrants that landed when weather was favorable for continued migration. Some of the variation in the energetic condition of arrivals may be explained by the location where migrants initiated crossings. Our simulation of flight over the Gulf of Mexico showed that with following winds a warbler can cross the Gulf of Mexico from Yucatan with fat reserves to spare, and stronger tailwinds make flights from as far south as Honduras energetically permissible. The length of stay after a trans-Gulf flight was related to the extent of fat-depletion upon arrival: lean birds stayed longer than fat migrants. Migrants stopped over for 1–7 days and replenished energy reserves at rates that varied from 0.19 g/d for Hooded Warblers (Wilsonia citrina) to 0.87 g/d for Ovenbirds (Seiurus aurocapillus). Within each species, most individuals gained weight at a rapid rate, though a few individuals lost or maintained weight during their stay.     

Avoidance of headwinds or exploitation of ground effect—why do birds fly low?

ABSTRACT Birds often fly close to the ground or water. Wind shear theory predicts that wind speeds decline with proximity to the substratum, so birds might be expected to fly lower when flying upwind than when flying downwind. We tested this prediction and found that the wind shear equation is valid at heights below 4 m, with wind speed over a smooth surface ～40% lower at a height of 0.08 m than at 4 m. Birds that fly close enough to smooth substrata can also benefit energetically from ground effect, where vortices generated by their flight interact with the ground or water. This suggests that birds should use ground effect more when flying upwind than when flying downwind. We determined the percent time spent flying in ground effect by 21 species of passerine and non‐passerine birds flying in sheltered coastal aquatic and nearby terrestrial areas of County Cork, Ireland. We found that use of ground effect was uncommon for passerines, but common for a variety of non‐passerine waterbirds. However, phylogenetic analysis indicates no linkage between phylogeny and incidence of ground effect use and it is probable that incidence of use is determined by ecology rather than phylogeny. Great Cormorants (Phalacrocorax carbo) used ground effect most frequently over water (59.4% of time in flight). Over land, Barn Swallows (Hirundo rustica) used ground effect most often (19.8% of time). Phylogenetic contrasts regression analysis showed no significant relationship between use of ground effect and either wing aspect ratio or wing loading for 18 of our focal species, though simple linear regression analysis indicated that birds with greater wing loading used ground effect slightly (but significantly) more often. We found that 95% of Great Cormorants flying upwind used ground effect whereas only 35% did so when flying downwind. Few Black‐headed Gulls (Chroicocephalus ridibundus) used ground effect (probably because they fly high to locate prey), but still showed greater use when flying upwind (25%) than downwind (2.5%). When flying upwind in ground effect at the wind speeds encountered in our study, the velocity for minimum power (V_mp) for Great Cormorants was exceeded, suggesting theoretical benefits of about 14.3%. Our study indicates that several species exploit both wind shear and ground effect to minimize energy expenditure during commuting and foraging, but that others do not, because of either complexity of habitat morphology or the demands of their foraging ecology.     

Seasonal differences in energy requirements of Garden Warblers Sylvia borin migrating across the Sahara desert

The Sahara desert acts as an ecological barrier for billions of passerine birds on their way to and from their African wintering areas. The Garden Warbler Sylvia borin is one of the most common migrants involved. We used body mass of this species from Greece in autumn and spring to simulate the desert crossing and to assess how body mass relates to fuel requirement. The flight range estimates were adjusted to the seasonal extent of the desert, 2200 km in autumn and about 2800 km in spring. In autumn, with an average fuel load of about 100% of body mass without fuel, birds were not able to cross the desert in still air, but northerly winds prevail during September and with the average wind assistance only one in 14 was predicted to fail to make the crossing. Body mass data from spring, after the desert crossing, was used to estimate departure body mass from south of the desert. The average wind assistance in spring is close to zero and departure body mass of the average bird arriving at Antikythira, a small Greek island, under such conditions was estimated to be 34.6 g, which corresponded to a fuel load of 116%. Calculations based on 1% body mass loss per hour of flight showed slightly larger body mass loss than that calculated from flight range estimates. The results suggest that passerine birds about to cross the eastern part of the Sahara desert need to attain a larger fuel load in spring than in autumn.     

Effects of moth size on velocity and steering during upwind flight toward a sex pheromone source byLymantria dispar (Lepidoptera: Lymantriidae)

Free-flying male gypsy moths (Lymantria dispar)head upwind in response to sex pheromone. Males typically fly in a zigzag path, with mean ground speeds modulated by pheromone concentration and ambient temperature, but not by wind speed. We studied the effect of male size on ground speeds and additional flight track parameters. Mean net ground speed along the wind line was fastest among large males and was slower in medium and small males. Similarly, mean airspeeds and ground speeds along the flight tracks increased from small to large males. Males from all three size classes steered similar mean course angles. Small males, however, had larger mean track angles than larger males, and mean drift angles were also larger for small males. Turning rates (frequency of turns across the wind line) and interturn distances (net crosswind displacement between turn apices) were not significantly different among the three size classes; however, large males had a trend toward a reduced mean turning rate and increased mean interturn distance. The steering of similar course angles by males from all three size classes and the higher airspeeds among larger males (the two variables males can actively control during free flight) suggest that changes in other flight parameters are a result primarily of increased ground speed among large males.