Nocturnal windborne migration of ground beetles,particularly Pseudoophonus griseus (Coleoptera: Carabidae), in China

1 A network of light‐traps, an aerial net carried by kytoon (balloon) and two entomological radars were used to investigate whether ground beetles migrate nocturnally through China. The network‐wide, simultaneous sudden increase in light trap catches, and after subsequent decrease, indicated a seasonal long‐distance night migration of ground beetles, with Pseudoophonus griseus (Panzer) predominant, in August. 2 Aerial net trapping indicated that carabids were able to ascend to altitudes of at least 200 m and become windborne. Radar observations indicated that the migratory beetles formed high‐density layer concentrations at approximately 200–300 m. 3 These concentrations were coincident with the top of the temperature inversion and a wind speed maximum, which suggested that the carabids tended to select warm, fast moving air for their long‐distance migration. 4 The ground beetles orientated and displaced towards the downwind direction in southerly winds. Their air speed decreased as the tailwind increased and, thus, migrating beetles appeared to be conserving energy. 5 The mean ± SD displacement speed (ground speed) and air speed were 6.85 ± 1.73 m/s (n= 172) and 4.45 ± 1.54 m/s (n= 172), respectively. The duration of flight, estimated from the variation in area density derived from radar data, was approximately 9–10 h, indicating that the beetles might migrate hundreds of kilometres in a single flight.     

Aphid aerial density profiles are consistent with turbulent advection amplifying flight behaviours: abandoning the epithet 'passive'

Seminal field studies led by C. G. Johnson in the 1940s and 1950s showed that aphid aerial density diminishes with height above the ground such that the linear regression coefficient, b, of log density on log height provides a single-parameter characterization of the vertical density profile. This coefficient decreases with increasing atmospheric stability, ranging from -0.27 for a fully convective boundary layer to -2.01 for a stable boundary layer. We combined a well-established Lagrangian stochastic model of atmospheric dispersal with simple models of aphid behaviour in order to account for the range of aerial density profiles. We show that these density distributions are consistent with the aphids producing just enough lift to become neutrally buoyant when they are in updraughts and ceasing to produce lift when they are in downdraughts. This active flight behaviour in a weak flier is thus distinctly different from the aerial dispersal of seeds and wingless arthropods, which is passive once these organisms have launched into the air. The novel findings from the model indicate that the epithet 'passive' often applied to the windborne migration of small winged insects is misleading and should be abandoned. The implications for the distances traversed by migrating aphids under various boundary-layer conditions are outlined.     

THE CHANGING NUMBERS OF APHIS FABAE SCOP., FLYING AT CROP LEVEL, IN RELATION TO CURRENT WEATHER AND TO THE POPULATION ON THE CROP

Aphis fabae Scop, flying over bean crops in the summer usually show a double peak of aerial density during the day; scarcely any flight occurs at night. The first peak is thought to be composed mainly of alatae moulted since the previous evening and the decline of the peak to their depletion as they fly away from the crop. The second peak is probably composed mainly of alatae moulted during the same day. Lack of flight at night is due partly to low temperature, partly to lack of alatae old enough to fly and probably also to low light intensity.
Contrary to expectation changes in aerial density from hour to hour are only very weakly correlated with weather factors, especially wind-speed. The total numbers in each of the two peaks do show a low but significant correlation with both wind-speed and temperature in one case, but even then only a relatively small amount of the variation in aerial numbers is associated with weather changes. Other factors, particularly rapid changes in the numbers of alatae on the crop caused by moulting, accumulation before flight and depletion by flight evidently obscure changes due simply to varying flight behaviour. Thus because of large populations on the crop, it is possible to have quite large numbers in the air even when the weather is relatively unfavourable for flight.
In addition to these observations, the relation between total numbers caught at different wind-speeds shows that most of the migration occurred in winds when the aphids could have had no control over the general direction of flight. The current view that most migration takes place only in calm weather can therefore no longer be held.     

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.     

Effects of age and mated status on flight potential of Helicoverpa armigera (Lepidoptera: Noctuidae)

Abstract The effects of ageing and female mated status on the flight potential of Helicoverpa armigera (Hübner), collected as larvae from a pigeonpea crop in southern India, were investigated using a tethered-flight technique. In non-mated moths fed sugar solution, from the first night after adult eclosion, the durations of both total and longest continuous flight per night increased up to night 4 and remained at this level until at least night 6. Ovarian maturation was rapid with 77% of unmated moths having commenced oviposition by the third night. On the basis of field evidence it is likely that most females would be mated by the third or fourth night, provided plants with nectar or sugary exudates were locally available. In successfully-mated females a 15-fold decrease in total flight duration and a 28-fold decrease in longest continuous flight duration was observed in contrast to non-mated females of similar age. As host plants suitable for adult feeding and oviposition were locally available during the time of feral adult emergence, synchronous pre-reproductive migration was unlikely to occur in the population studied.     

High-Altitude Windborne Transport of <Emphasis Type="BoldItalic">Helicoverpa armigera</Emphasis> (Lepidoptera: Noctuidae) in Mid-Summer in Northern China

The high-altitude windborne transport of the second-generation moths of Helicoverpa armigera (Hübner) was observed with radar in July 2001 and 2002 at Langfang, Hebei province, China. Moths of local populations took off at dusk, with area density peaking ca. 0.5 h later. The variation of area density through the night suggested that the maximum flight duration was about 8.5 h. The moths generally ascended to altitudes of up to 1500 m above ground level (AGL) and formed layer concentrations at ca. 300, 500, and 1000 m AGL. Multiple layering was very common and layers were located at zones of maxima in wind speed or in wind shear rather than in temperature inversions. Both the displacement direction and the orientation of the insect were at an acute angle to the downwind direction, rather than being distributed at random, and varied as the wind direction changed. The windborne moths were mainly transported into northeastern China where maize at the silking stage would have provided good hosts for the subsequent (third) generation of H. armigera. These population movements may account for the nondevelopment of resistance to Bt cotton in H. armigera in northern China.     

THE FEASIBILITY OF USING VERTICAL-LOOKING RADAR TO MONITOR THE MIGRATION OF BROWN PLANTHOPPER AND OTHER INSECT PESTS OF RICE IN CHINA

Abstract The recent development of automatically operating, inexpensive vertical-looking radar (VLR) for entomological purposes has made it practical to carry out routine, automated monitoring of insect aerial migration throughout the year. In this paper we investigate whether such radars might have a role in monitoring and forecasting schemes designed to improve the management of the Brown Planthopper (BPH), Nilaparvata lugens , and of associated rice pest species in China. A survey of the literature revealed that these insects typically migrate at altitudes between 300 to 2 000 m above ground level, but calculations based on BPH radar scattering cross-sections indicated that the maximum altitude at which they individually produce signals analysable by current VLRs is only ˜ 240 m. We also show that coverage over most of the flight altitudes of BPH could be achieved by building a VLR using a wavelength of 8.8 mm instead of the 3.2 cm of existing VLR, but that such a radar would be expensive to build and to operate. We suggest that a more practical solution would be to use a 3.2 cm VLR as a monitor of the aerial movement of the larger species, from which the migration of rice pests in general might be inferred.     

Flight duration of the brown planthopper, Nilaparvata lugens (Homoptera: Delphacidae)

ABSTRACT.

• 1 The brown planthopper, Nilaparvata lugens (Stål), is a major pest of rice in Asia. It is known to make wind-assisted migratory flights each year to colonize the summer rice growing areas of China, Japan and Korea.
• 2 Modelling windborne displacements between rice growing areas in Asia requires migratory behaviour and flight duration to be established for this insect.
• 3 Field and laboratory observations suggest that N. lugens take-off at dusk and that some continue flying for up to 24–26 h if the temperature is ≥ 17°C.
• 4 Trajectories for 10 m above ground level and 1.5 km above mean sea level are used to identify possible sources and, hence, to estimate the flight times of N.lugens caught in nets on ships on the East China Sea in 1973 and 1981.
• 5 Estimated flight times between the sources and the ships ranged from about 9 to 30 h.
• 6 Results suggest that long-distance migration can occur in surface winds, when they are strong, but that long-distance migration is more likely at 1.5 km.
• 7 When simulating windborne displacements of N.lugens, it can be assumed that in areas and at heights where the temperature is ≥ 17°C, some migrants will fly downwind for up to 30 h after a dusk take-off. Others will fly for shorter periods, giving the population as a whole the opportunity to colonize all the rice crops flown over.

     

Strategies of passerine migration across the Mediterranean Sea and the Sahara Desert: a radar study

Radar observations of the diurnal timing of bird migration in the Sahara Desert are presented for autumn migration. Study sites were on a transect along the north-south migratory direction. Three groups of birds migrating either during day, evening or night in the northern part of the Western desert in Egypt were identified. The maximum of day and night groups occurred later the further south the study sites were. Based on the distance between sites and the timing of peak migration, birds were flying at an estimated ground speed of about 20 m/s. The maximum of the evening group was at about 21:00 h at all sites. The three groups were classified according to three different strategies of migration across the Mediterranean Sea and the Sahara Desert: (1) the day group of birds performed a non-stop flight across the sea and at least the northern part of the desert; [2] the night group performed an intermittent migratory strategy with stopover at the coast of Egypt to continue migration the next evening; (3) the evening group birds were also intermittent migratory fliers, but they stopped somewhere in the desert after a continuous flight across the sea and part of the desert. About 20% of all migrants are involved in non-stop migration and 80% in intermittent migration with stopover at the coast (70%) or with stopover in the desert (10%). It is argued that any species of small passerine has the option to use any of the three strategies.     

Variation in the nocturnal flight behaviour of migratory birds along the northwest coast of the Mediterranean Sea

The Mediterranean Sea is one of the largest obstacles that has to be crossed by Palearctic birds migrating from Europe to Africa; it thus offers a good opportunity to study variations in migratory behaviour of birds facing a major ecological barrier. Using a passive infrared device, the flight directions of nocturnal migrants were determined and flight altitudes estimated at ten sites along the French and Spanish coast of the Mediterranean Sea in September and October 1995. The variation of migratory intensity, flight direction and altitude in the course of the night was examined. The highest density of migration was recorded within the first hour after sunset, followed by relatively high densities over the next several hours, and a progressive decrease in the last third of the night. In spite of broad variation in the course of the coastline relative to the basic directions of migration and specific reactions of the migrants to the local conditions, a decrease in seaward migration corresponding to an increase in landward migration from the first to the second half of the night was a general feature at nearly all sites. The results suggest a shift in the motivation of the birds depending on the time of arrival in a coastal area, leading to an adjustment in the flight behaviour of nocturnal migrants.     

Diel vertical movements,and effects of infection by the cestode <Emphasis Type="Italic">Schistocephalus solidus</Emphasis> on daytime proximity of three-spined sticklebacks <Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis> to the surface of a large Alaskan lake

We conducted a field study in Iliamna Lake, Alaska, to test the hypothesis that proximity of three-spined sticklebacks Gasterosteus aculeatus to the lake’s surface during the daytime varies with macroparasitic cestode parasite Schistocephalus solidus infection in a manner consistent with enhanced vulnerability to avian predators. Extensive sampling in the lake and likelihood-based modeling revealed that sticklebacks displayed a diel vertical migration, being closer to the surface at night than during the evening and early morning. Additional sampling, also coupled with a likelihood-based modeling approach, showed that fish caught at the surface of the lake during the day were more often parasitized (76 vs. 65%), more heavily parasitized (26.8 vs. 22.7% of their body mass), and had larger individual parasites (0.24 vs. 0.20 g) than those caught at night. Parasite infection was related, non-linearly, to fish size, which also differed between day and night sampling at the surface. We performed statistical competitions among nested hierarchies of models that accounted for this effect of length. The most likely models indicated that fish captured during the day had greater parasite prevalence, higher parasite burdens, and larger parasites than did fish captured at night. Proximity to the surface during the day in this very clear lake would likely increase the vulnerability of sticklebacks to predation from birds, enabling completion of the parasite’s lifecycle.     

Aerial distribution, flight behaviour and eggload: their inter-relationship during dispersal by the sweetpotato whitefly

1. The aerial distribution of Bemisia tabaci Gennadius (the sweetpotato whitefly) was studied during the early ascent phase of flight, to test the degree to which dispersal patterns reflect the flight behaviour of individuals.
2. Marked whiteflies were trapped at four heights between 0 and 7·2 m above fallow ground, and at six distances between 0 and 100 m from the insect source. Insects were trapped during a 2–3 h period after the initiation of flight activity during the summers of 1995 and 1996.
3. Analysis of trap catch data revealed a clear negative exponential relationship between height and aerial distribution, and a slightly weaker negative power relationship between distance and aerial distribution. Marked insects were caught in the uppermost traps adjacent to the source, indicating that a portion of the population had a strong capacity for ascent out of the flight boundary layer.
4. Eggload decreased with the height, but not the distance, at which whiteflies were trapped. Mean eggload close to the ground was significantly greater than that for those trapped at 4·8 and 7·2 m, supporting the hypothesis that there is a trade-off between flight and oogenesis in weak-flying insects.
5. Air temperatures during the trapping periods were positively correlated with the proportion of male and female B. tabaci caught in the highest traps, but not in the most distant traps.
6. The significance of these results for accurate prediction of whitefly dispersal is discussed, and the importance of individual's behaviour in determining dispersal patterns of small insects is emphasized.     

Environmental effects on flying migrants revealed by radar

Migratory animals are affected by various factors during their journeys, and the study of animal movement by radars has been instrumental in revealing key influences of the environment on flying migrants. Radars enable the simultaneous tracking of many individuals of almost all sizes within the radar range during day and night, and under low visibility conditions. We review how atmospheric conditions, geographic features and human development affect the behavior of migrating insects and birds as recorded by radars. We focus on flight initiation and termination, as well as in‐flight behavior that includes changes in animal flight direction, speed and altitude. We have identified several similarities and differences in the behavioral responses of aerial migrants including an overlooked similarity in the use of thermal updrafts by very small (e.g. aphids) and very large (e.g. vultures) migrants. We propose that many aerial migrants modulate their migratory flights in relation to the interaction between atmospheric conditions and geographic features. For example, aerial migrants that encounter crosswind may terminate their flight or continue their migration and may also drift or compensate for lateral displacement depending on their position (over land, near the coast or over sea). We propose several promising directions for future research, including the development and application of algorithms for tracking insects, bats and large aggregations of animals using weather radars. Additionally, an important contribution will be the spatial expansion of aeroecological radar studies to Africa, most of Asia and South America where no such studies have been undertaken. Quantifying the role of migrants in ecosystems and specifically estimating the number of departing birds from stopover sites using low‐elevation radar scans is important for quantifying migrant–habitat relationships. This information, together with estimates of population demographics and migrant abundance, can help resolve the long‐term dynamics of migrant populations facing large‐scale environmental changes.     

Wingtip shape and flight performance in the European Starling Sturnus vulgaris

Although wingform is known to differ among individuals of the same species it is not known how intraspecific variation in wingtip shape is associated with flight performance. In this study, we have examined both among- and within-individual variation in wingtip shape in relation to changes in flight performance in the European Starling Sturnus vulgaris . We found that level flapping-flight speed and the ability to negotiate an aerial obstacle course were unrelated to wingtip shape. However, take-off parameters did vary with wingtip shape; birds with more rounded wingtips tended to take off from the ground at a steeper angle of ascent than those with relatively more pointed wingtips. The same relationships between wingtip morphology and flight were present in both the inter- and intra-individual experimental analyses. The evolutionary importance of this variation in take-off ability is discussed in terms of predator avoidance and enhancement of individual survival.     

Radar observations of the spring migration into northeastern China of the oriental armyworm moth, Mythimna separata, and other insects

Abstract. 1. The spring migration of the oriental armyworm moth, Mythimna separata (Walker), and other insects into northeastern China was observed by radar at a site in central Jilin province. Samples of the migrants were obtained in a net flown from a kite, and M.separata populations in the surrounding region were monitored with a trap network. 2. The radar regularly detected echoes which were of the type characteristic of large insects, and three out of four large insects in the aerial samples were noctuid moths, including one M.separata. Catches of armyworm moths in the regional trap network peaked during the period of radar observations. 3. Migration occurred at night. It commenced with a take-off flight at dusk and generally continued until dawn, with numbers often being highest around midnight. Most migration took place at altitudes below 500 m, with strong layer concentrations forming at 200–400 m during the middle part of some nights. 4. Migration was approximately downwind. The net movement was overwhelmingly to the northeast because southwesterly winds occurred most frequently and were relatively strong, and because migration was more intense and prolonged on these winds. Orientation tended to be to the left of the downwind direction, and was most often to the north or northeast. 5. Migration into northeastern China was accomplished in a series of night-time movements of 100–300 km rather than by a single non-stop flight. 6. The net movement of insects towards the northeast was sufficient to produce the observed regional infestation of M.separata moths. Oviposition by immigrant armyworm moths in vulnerable crops would have been at a level where economically significant damage would be expected from the resulting larval population. 7. Analogous springtime migrations of noctuids leading to temporary colonizations of habitats at higher latitudes occur in other continents. Such colonizations may be unproductive if, as appears possibly the case for northeastern China, prevailing winds later in the season are generally unfavourable for a return migration towards overwintering areas at lower latitudes.     

The bio-rhythmic problems of a space flight

The influence of space-flight factors on the organism's circadian periodicity is discussed. It is shown that in mechanisms of such influence, the transformation of natural structure of Zeitgebers accompanying a space flight, can play the essential role. It is confirmed by the results of ground research carried out in conditions of isolation simulating the usual space flight transformation of Zeitgebers system. The data obtained in the ground researches with isolation, testify that the changes of circadian rhythms in these conditions are similar to those in space flights and frequently are interpreted as the result of influence of weightlessness. Special attention is paid to phenomenon of delay of the daily maximum and to possible connection of this phenomenon with the tendency to transition of rhythmic process in a free-running regime.     

Downwind migration of the African army worm moth, Spodoptera exempta, studied by mark-and-capture and by radar

Abstract. 1. About 166,000 African armyworm moths, Spodoptera exempta (Walk.), were marked at an emergence site near Nairobi when they fed at night on trees baited with dyed molasses.
2. Six marked moths were captured in pheromone traps, including one at 90 km after flying for only one night, and another at 147 km.
3. Moth flight trajectories deduced from radar and from marking showed that migration was downwind.
4. During migration, moths become dispersed; hence the high densities that lead to outbreaks must be produced by concentration.
5. Some moths were ready to mate on the same night they completed their long-distance flight.     

Wind selection and drift compensation optimize migratory pathways in a high-flying moth

Numerous insect species undertake regular seasonal migrations in order to exploit temporary breeding habitats [1]. These migrations are often achieved by high-altitude windborne movement at night [2-6], facilitating rapid long-distance transport, but seemingly at the cost of frequent displacement in highly disadvantageous directions (the so-called "pied piper" phenomenon [7]). This has lead to uncertainty about the mechanisms migrant insects use to control their migratory directions [8, 9]. Here we show that, far from being at the mercy of the wind, nocturnal moths have unexpectedly complex behavioral mechanisms that guide their migratory flight paths in seasonally-favorable directions. Using entomological radar, we demonstrate that free-flying individuals of the migratory noctuid moth Autographa gamma actively select fast, high-altitude airstreams moving in a direction that is highly beneficial for their autumn migration. They also exhibit common orientation close to the downwind direction, thus maximizing the rectilinear distance traveled. Most unexpectedly, we find that when winds are not closely aligned with the moth's preferred heading (toward the SSW), they compensate for cross-wind drift, thus increasing the probability of reaching their overwintering range. We conclude that nocturnally migrating moths use a compass and an inherited preferred direction to optimize their migratory track.     

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.     

Return migration of Helicoverpa armigera (Lepidoptera: Noctuidae) during autumn in northern China

The autumn migration of Helicoverpa armigera (Hübner) was observed with radar and two types of light-trap at Langfang, Hebei province, China in 2001 and 2002. The sudden increase in the proportion of H. armigera moths in the searchlight trap indicated migration into the area and catches increased 10-fold during the second half of the night due to the landing of migrants before dawn. The moths' migratory flights took place at up to 2000 m above the ground, and moths flew differentially at times, and heights, when favourable (i.e. northerly) winds occurred. This facilitated the maximum displacement of moths towards the south during these 'return' migrations. The moths flew over the radar site at consistently high densities through the night, and the resulting flight durations of c. 10 h, at displacement speeds of 30-33 km h-1, would allow moths emerging in the far northeast of China (i.e. Liaoning and Jilin provinces and the Inner Mongolia autonomous region) to migrate into northern China (Hebei, Shandong and Henan provinces). The association of the seasonal migratory movements of H. armigera with crops in northern China is briefly discussed.