Modeling distributions of flying insects: effective attraction radius of pheromone in two and three dimensions

The effective attraction radius (EAR) of an attractive pheromone-baited trap was defined as the radius of a passive “sticky” sphere that would intercept the same number of flying insects as the attractant. The EAR for a particular attractant and insect species in nature is easily determined by a catch ratio on attractive and passive (unbaited) traps, and the interception area of the passive trap. The spherical EAR can be transformed into a circular EAR_c that is convenient to use in two-dimensional encounter rate models of mass trapping and mating disruption with semiochemicals to control insects. The EAR_c equation requires an estimate of the effective thickness of the layer where the insect flies in search of mates and food/habitat. The standard deviation (SD) of flight height of several insect species was determined from their catches on traps of increasing heights reported in the literature. The thickness of the effective flight layer (F_L) was assumed to be , because the probability area equal to the height of the normal distribution,, times the F_L is equal to the area under the normal curve. To test this assumption, 2000 simulated insects were allowed to fly in a three-dimensional correlated random walk in a 10-m thick layer where an algorithm caused them to redistribute according to a normal distribution with specified SD and mean at the midpoint of this layer. Under the same conditions, a spherical EAR was placed at the center of the 10-m layer and intercepted flying insects distributed normally for a set period. The number caught was equivalent to that caught in another simulation with a uniform flight density in a narrower layer equal to F_L, thus verifying the equation to calculate F_L. The EAR and F_L were used to obtain a smaller EAR_c for use in a two-dimensional model that caught an equivalent number of insects as that with EAR in three dimensions. This verifies that the F_L estimation equation and EAR to EAR_c conversion methods are appropriate.     

An evaluation of western bean cutworm pheromone trapping techniques (Lepidoptera: Noctuidae) in a corn and soybean agroecosystem

Pheromone traps can be used to monitor for adult western bean cutworms, Striacosta albicosta (Smith) (Lepidoptera: Noctuidae), and for the timing of field scouting. Understanding the effect that different trapping techniques have on adult captures could help corn (Zea mays L.) producers make better pest management decisions. Several approaches to trapping adults were evaluated in 2005 and 2006 by using two different pheromone traps (sticky wing and jug traps) in two different environments (corn or corn/soybean [Glycine max (L.) Merr.] at three different heights (0.6, 1.2, and 1.8 m). There was no significant difference in the trap catches by trap type in either 2005 or 2006. There were significantly more adults captured in traps placed between two cornfields than traps placed between corn/soybean fields during both years. Trap height also was significant, with the traps at 1.2 and 1.8 m catching more moths than traps at 0.6 m during both years. These results show that trapping techniques do affect trap catches and that either trap type placed between two cornfields at either 1.2 or 1.8 m above the ground will maximize trap catches.     

Stable fly (Stomoxys calcitrans: Diptera, Muscidae) trap response to changes in effective trap height caused by growing vegetation

Stable flies (Stomoxys calcitrans L.) are blood-feeding pests of cattle, whose populations are often monitored using sticky traps. Trap responses at different heights were compared with: 1) a choice and no-choice test, with sticky cards set at 30 and 121 cm heights (above ground), and 2) tall (120 cm) traps placed in short (3 cm) and tall (40 cm) grass to assess how vegetation height affects trap catches. In the first experiment, the percentage of upper to lower trap catches were similar at choice (16%) and no-choice traps (15%). In the vegetation study, stable fly catch height data were fitted to gamma distributions to determine the most productive trap heights; 20 cm above short grass and 24 cm above tall grass (from lower edge of trap). The results indicate that traps used to monitor stable fly populations should be maintained at a constant distance above surrounding vegetation rather than ground surface, otherwise trap data can be misleading.     

Traps and attractants for wood-boring insects in ponderosa pine stands in the Black Hills, South Dakota

Recent large-scale wildfires have increased populations of wood-boring insects in the Black Hills of South Dakota. Because little is known about possible impacts of wood-boring insects in the Black Hills, land managers are interested in developing monitoring techniques such as flight trapping with semiochemical baits. Two trap designs and four semiochemical attractants were tested in a recently burned ponderosa pine, Pinus ponderosa Dougl. ex Laws., forest in the Black Hills. Modified panel and funnel traps were tested in combination with the attractants, which included a woodborer standard (ethanol and alpha-pinene), standard plus 3-carene, standard plus ipsenol, and standard plus ipsdienol. We found that funnel traps were equally efficient or more efficient in capturing wood-boring insects than modified panel traps. Trap catches of cerambycids increased when we added the Ips spp. pheromone components (ipsenol or ipsdienol) or the host monoterpene (3-carene) to the woodborer standard. During the summers of 2003 and 2004, 18 cerambycid, 14 buprestid, and five siricid species were collected. One species of cerambycid, Monochamus clamator (LeConte), composed 49 and 40% of the 2003 and 2004 trap catches, respectively. Two other cerambycids, Acanthocinus obliquus (LeConte) and Acmaeops proteus (Kirby), also were frequently collected. Flight trap data indicated that some species were present throughout the summer, whereas others were caught only at the beginning or end of the summer.     

Measuring trap efficiency for bark beetles (Col., Scolytidae)

Abstract:  The relative efficiency of cylindrical, linear and cross-barrier traps for trapping bark beetles was investigated based on a theoretical model. Using this model, the effective trap interception area of each trap type was calculated and trap efficiency was defined as the ratio of the effective interception area to the trap surface area. The relative efficiencies of the three trap types were calculated as the ratios of their respective effective interception areas. Based on this approach, assuming random directional movement of dispersing beetles, the order of efficiency of the three trap types, from highest to lowest, was linear, cross-barrier and cylindrical. The expected ratios of trap catches based on the relative efficiencies of the three trap types were fitted to data from trapping experiments with the mountain pine beetle ( Dendroctonus ponderosae Hopkins). In general, there was large variation in trap catches among traps of the same type but the ratios of mean catches per trap conformed to the expected ratios. The results indicate that the model of trap efficiency could be used for designing efficient traps. The methods presented are amenable for assessing the efficiency of other trap designs.     

Uraba lugens (Lepidoptera: Nolidae) in New Zealand: pheromone trapping for delimitation and phenology

A synthetic sex pheromone trapping survey of the leaf skeletonizer Uraba lugens Walker (Lepidoptera: Nolidae) demonstrated the unexpectedly widespread distribution of the insect across > 40,000 ha of urban Auckland, New Zealand. A survey of eucalyptus trees planted in parks and other public areas showed a significant spatial correlation between trap catch and breeding populations, validating the trap survey results. Traps in trees showing damage had four-fold higher catches than traps placed in undamaged or nonhost trees, and < 1% of damaged trees with traps failed to catch adult moths. Damage by larval feeding was correlated with male trap catch in the previous generation, offering good prospects for a pest management decision support system, provided that an economic threshold is developed. Catches increased by 3.4-fold in the same georeferenced trapping grid between November and December 2003 and between March and April 2004 across two generations, over the summer. A vertical transect showed that catches increased with height up to the top trap at 13 m (60% of mean tree height). Options for managing the insect will need to overcome the high rate of increase, the rate of spread, and the vertical distribution of the insect on tall eucalyptus trees.     

Optimising coloured pan traps to survey flower visiting insects

Colour is an important attractant for many flower-visiting insects (anthophiles). Consequently, coloured pan trapping is an efficient technique that can be easily and cost-effectively used to quantitatively sample assemblages of anthophiles. However, colour preferences of anthophiles is an important source of bias that needs to be considered in pan trap surveys. By drawing sub samples comprised of different colour combinations from a database of pan trap surveys in the lowlands of the Cape Floristic Region, we examine the effects of colour on pan trap catches and determine which combinations of colours might provide better estimates of diversity when sampling with multi-colour sets of pan traps. Pan trap catches included the major groups of anthophiles in the region (Coleoptera, Diptera and Hymenoptera), but butterflies (Lepidoptera) were strongly under-represented. Colour played an important role in determining the species richness and composition of pan trap catches, with colour sets that included high reflectance yellow and white generally having catches with the highest species richness. While all colour combinations provided reasonable estimates of proportional species richness, proportional abundance of taxa varied among different colour sets, and did not accurately reflect actual proportions of different taxa in the entire dataset. For comparative biodiversity surveys and assessments we recommend the use of high reflectance colours such as white and yellow traps, while for full inventory surveys, it may be necessary to include other colours to catch rarer species that are excluded by the high reflectance colours alone.     

Efficacy of CC traps and seasonal activity of adult Bemisia argentifolii (Homoptera: Aleyrodidae) in Imperial and Palo Verde Valleys, California

Adult whitefly Bemisia argentifolii Bellows & Perring trap (CC trap) catches were compared with suction type trap catches. CC trap catches were significantly correlated to the suction trap catches. Higher numbers of B. argentifolii adults were caught in CC traps oriented toward an untreated, B. argentifolii-infested, cotton field as compared with traps oriented toward Bermuda grass fields, farm roads, or fallow areas. CC trap catches at five heights above ground (from 0 to 120 cm) were significantly related to each other in choice and no-choice studies. CC trap catches were low in the Imperial and Palo Verde Valleys from late October to early June each of 1996, 1997, and 1998. Trap catches increased with increasing seasonal air temperatures and host availability. Trap catches were adversely affected by wind and rain. Abrupt trap catch increases of 40- to 50-fold for 1-2 d in late June to early July followed by abrupt decreases in adult catches suggest migrating activity of adults from other nearby crop sources.     

Efficiency of Malaise traps and colored pan traps for collecting flower visiting insects from three forested ecosystems

Pan and Malaise traps have been used widely to sample insect abundance and diversity, but no studies have compared their performance for sampling pollinators in forested ecosystems. Malaise trap design and color of pan traps are important parameters that influence insect pollinator catches. We compared pan trap (blue, yellow, white, and red) and Malaise trap catches from forests in three physiographic provinces (Piedmont, Coastal Plain, and Blue Ridge) of the southeastern United States. Similarities in trap performance between sites were observed with blue pan traps being most effective overall. Our results showed that various pollinator groups preferred certain pan trap colors and that adding color to Malaise traps influenced insect pollinator catches. However, pan traps generally caught more pollinators than Malaise traps. Because of their low cost and simplicity, using several colors of pan traps is an effective way to sample relative abundance and species richness of flower-visiting insects.     

Effect of height and color on the efficiency of pole traps for Aethina tumida (Coleoptera: Nitidulidae)

Olfactory cues released by adult bees, brood, pollen, and honey from a honey bee, Apis mellifera L., colony are the primary stimuli that guide the beetle Aethina tumida Murray (Coleoptera: Nitidulidae) to host colonies. To investigate the response of adult A. tumida to visual stimuli, we tested the influence of color and height on trap efficiency. Two pole trap colors (black and white) were evaluated at three heights (46 cm, 1 m, and 3m) from October 2008 to December 2009. A. tumida were trapped in the greatest numbers between 17 April and 15 May 2009. The lowest numbers were captured during the winter and fall. The trapping results showed that both color and trap height significantly influenced capture. The average catch in the white traps (mean +/- SE, 2.47 +/- 0.30) was significantly higher than that of the black traps (1.53 +/- 0.29) probably because white is more reflective than black. Among the heights evaluated, there were more beetles caught when traps were positioned at 46 cm (the same height as the entrance of the hives) with 3.07 +/- 0.51 beetles compared with beetles captured at 1 m (1.88 +/- 0.30) or 3 m (1.06 +/- 0.18) high. Male and female beetles exhibited similar responses to trap color and height. The relationship between the numbers of beetles in colonies and capture rates in traps was very poor and did not provide a basis to evaluate trap efficiency. In addition, because capture rates seemed generally low in relationship to the number of beetles in the apiary, substantial improvements to the trap may be necessary.     

Observations on different methods of aphid trapping

Cylindrical and horizontal sticky traps painted in a range of spectral colours were used to determine the flight and landing behaviour of aphids. Data are also presented on aphid catches in suction traps at two heights and in light traps. Apart from colour sensitivity (yellow versus white) there was apparently a separate response to colour which in some species varied with season. Within yellow-sensitive species there was also a differential response to colour. With the experimental methods used, it was not possible to define mathematically the active and passive landing components on cylindrical traps. Though the active landing component was large it varied between aphid species. Most species caught on horizontal traps at ground level had been flying above 1 m. In some species the response of males to colour and their landing behaviour differed from that of viviparae and oviparae. Four years data from suction traps suggest that aphid species can be divided into three categories on the basis of the height at which they normally fly. One group, mostly tree-feeders, always show the greatest density at a high level (12.2 m) throughout the season. The second group always have the highest density at a low level (1 m) whilst a third group of species change at a specific date each autumn from a maximum density at 1–12.2 m. Attraction to light (moth trap) appeared to be linked with the grouping of species by height of flight. The interpretation of catch data is discussed in the light of these observations.     

Visual Responses of Adult Asian Citrus Psyllid (Hemiptera: Liviidae) to Colored Sticky Traps on Citrus Trees

The effects of five differently-colored sticky traps in capturing adult Diaphorina citri were evaluated in citrus orchards. Trap catches of D. citri were monitored fortnightly on blue, green, red, white and yellow sticky cards placed on three citrus varieties during D. citri active flight period from April to July in south Texas. Evaluation of mean trap catches of each color by repeated measures analysis of variance produced three separate groups: yellow traps caught significantly more D. citri adults than the other four traps; red and green traps caught significantly more D. citri than blue and white traps, which were not significantly different. Although the number of adult psyllid captured on all trap types significantly increased with time during the trapping period, the performance of traps did not change with time. Trap catches were also significantly influenced by the citrus species; traps placed on lemon trees captured more D. citri than those placed on sweet orange and grapefruit, suggesting that plant preference exhibited by D. citri may influence the performance of traps. The ratio of trap reflectance between the 680 to 700 nm and the 450 nm was significantly correlated with total trap catches in all host species studied. Thus, this index was a good indicator of the attractiveness of adult D. citri to colored traps. Additionally, we compared the reflectance values of young versus mature flush shoots of the three host plants used in this study as related to densities of D. citri recorded in colored traps. We discussed the importance of visual cues in the host finding behavior of adult D. citri.     

Small‐plot studies comparing pheromone and juice baits for mass‐trapping invasive Synanthedon myopaeformis in Canada

Recent introduction of Synanthedon myopaeformis (Borkhausen) (Lepidoptera: Sesiidae) into organic apple‐growing areas of Canada has stimulated research on semiochemical‐based management of this European pest. Replicated, small‐plot (0.16 ha) experiments were conducted to compare sex pheromone, 3Z,13Z‐octadecadienyl acetate (10 mg), Concord grape juice (300 ml), or their combination, as mass‐trapping lures at trap densities equivalent to 12.5, 25, 50, and 100 traps ha^?1. Total numbers of male and female moths removed from test plots increased significantly with trap density in all juice‐based mass‐trapping experiments. In pheromone mass‐trapping experiments, however, total catches of males did not increase significantly as trap densities were increased and catches appeared to plateau with 25–50 traps ha^?1. With pheromone‐based mass‐trapping, significantly fewer males were caught in pheromone‐baited assessment traps at the centre of each mass‐trapping plot than in identical traps in untreated plots. This reduction is indicative of significant trap interference or trap ‘shut‐down’. Increasing the density of juice‐based mass‐trapping had no effect on catches of male or female moths in juice‐baited assessment traps, indicating a short range of attraction and lack of interference between juice traps. Pheromone‐ and juice‐based mass trapping removed similar numbers of males at each trap density tested, respectively, but summed catches of males and females were greatest with juice baits. Combining pheromone and juice into a single mass‐trapping treatment (50 traps ha^?1) did not significantly increase catches of males or females relative to either treatment alone. If a practical bisexual mass‐trapping system is going to be developed for S. myopaeformis, then identification of volatile kairomones in Concord grape juice may be useful.     

Factors affecting male Prays oleae (Lepidoptera: Yponomeutidae) captures in pheromone-baited traps in olive orchards

Effects of trap design, height and side of trap placement on olive trees, pheromone doses in dispensers, aging of dispensers in the field, and secondary pheromone components were evaluated for the development of an effective pheromone monitoring system for Prays oleae (Bernard) Lesne (Lepidoptera: Yponomeutidae) in olive orchards. Field trials showed that trap design, pheromone dose, and trapping side affected male captures, whereas dispenser age, trap height, and secondary components had no influence. Pherocon 1C and Delta traps baited with 1 mg of (Z)-7-tetradecenal captured more male moths than Pherocon II or Funnel traps. Placement of traps at different cardinal directions significantly affected captures, but this trend was not consistent and varied with flight period and trap position internal or external to the tree canopy. Moth phenology as determined by pheromone traps from early April to mid-October was consistent with published field data. Results indicate that Pherocon 1C or Delta traps baited with 1 mg of (Z)-7-tetradecenal provide an effective tool for monitoring the flight activity of P. oleae and the time of application of control measures.     

Trap type, chirality of alpha-pinene, and geographic region affect sampling efficiency of root and lower stem insects in pine.

Root and lower stem insects cause significant damage to conifers, vector phytopathogenic fungi, and can predispose trees to bark beetle attacks. The development of effective sampling techniques is an important component in managing these cryptic insects. We tested the effects of trap type and stereochemistry of alpha-pinene, in combination with ethanol, on catches of the root colonizing weevils (Coleoptera: Curculionidae) Hylobius spp. [mostly Hylobius pales (Herbst)], and Pachylobius picivorus (Germar), the root colonizing bark beetle (Coleoptera: Scolytidae) Hylastes porculus Erickson, and the lower stem colonizing bark beetle Dendroctonus valens (LeConte). We tested for inter-regional differences by conducting similar field assays in the northern (Wisconsin) and southern (Louisiana) United States. The more effective trap type varied with region. Root weevils were caught primarily in pitfall traps in Wisconsin, whereas they were caught mostly in lower stem flight traps in Louisiana. In Wisconsin, root colonizing bark beetles were also caught primarily in pitfall traps, but lower stem colonizing bark beetles were caught primarily in lower stem flight traps. The root feeding weevils preferred (-) over (+)-alpha-pinene in both regions. Some exceptions relating to trap type or gender occurred in southern populations. The two root and lower stem colonizing bark beetles in Wisconsin showed no preference between (+) and (-)-alpha-pinene in combination with ethanol. No bark beetles were caught in the south. Our results suggest that modifying trap type and enantiomeric ratios of monoterpenes for different insect groups and in different regions can improve sampling efficiency for these important pests.     

Distribution and dynamics of Japanese beetles along the Indianapolis Airport perimeter and the influence of land use on trap catch

As part of a program to minimize the accidental transportation of Japanese beetles (Popillia japonica) through cargo aircraft to areas where they are not established, a 4-yr trapping project was initiated to study the relative distribution and dynamics of the beetles along a trap line around the Indianapolis International Airport. Land use influence on beetle abundance (trap catch) was assessed using a geographic information system. Trap catch was consistently high in some locations and low in others. In general, high trap catches occurred near agronomic land planted with corn or soybeans, which are both preferred hosts of adult beetles. Low trap catches generally occurred in areas lacking preferred host plants. The amount of agronomic land within 500 m of the traps was always positively correlated with trap catch. Average trap catches were highly correlated by location from year to year, indicating stability of the relative distribution of the beetles along the trap line. Because high trap catches consistently occurred in the same locations, it can be inferred that trapping can be an effective method to monitor Japanese beetle populations. Taking airport-owned agronomic land out of corn and soybean production near the cargo terminals may reduce beetle activity in these areas.     

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.     

Female sex pheromone of brinjal fruit and shoot borer,Leucinodes orbonalis (Lepidoptera: Pyralidae): trap optimization and application in IPM trials

Delta and wing traps baited with synthetic female sex pheromone of Leucinodes orbonalis Guenée were found to catch and retain ten times more moths than either Spodoptera or uni-trap designs. Locally-produced water and funnel traps were as effective as delta traps, although 'windows' cut in the side panels of delta traps significantly increased trap catch from 0.4 to 2.3 moths per trap per night. Trap catch was found to be proportional to the radius of sticky disc traps in the range 5-20 cm radius, discs with a 2.5 cm radius caught no moths. Wing traps placed at crop height caught significantly more moths than traps placed 0.5 m above or below the crop canopy. Replicated integrated pest management (IPM) trials (3 x 0.5 ha per treatment) were conducted in farmers fields with young and mature eggplant crops. Farmers applied insecticides at least three times a week in all check and IPM plots. In addition pheromone traps were placed out at a density of 100 per ha and infested shoots removed weekly in the 0.5 ha IPM plots. Pheromone trap catches were reduced significantly from 2.0 to 0.4 moths per trap per night respectively in check and IPM plots in a young crop and 1.1 to 0.3 moths per trap per night in check and IPM plots respectively in a mature crop. Fruit damage was significantly reduced from an average of 41.8% and 51.2% in check plots of young and mature crops respectively to 22% and 26.4 respectively in the associated IPM plots. Significant differences in pheromone trap catches and fruit damage were attained four and two weeks respectively after IPM treatments began in the mature crop whereas in the immature crop significant differences were not observed for the first eight to nine weeks respectively. The relative impact of removing infested shoots and mass trapping on L. orbonalis larval populations was not established in these trials but in both cases there was an estimated increase of approximately 50% in marketable fruit obtained by the combination of control techniques compared to insecticide treatment alone.     

Aphis migration and the efficiency of the trapping method

Reproducible results were obtained when adhesive traps for aphides were placed at the same height in different parts of a potato field. The total catches in different localities in different seasons can therefore justifiably be compared. During the summer dispersal flights, different species flew at different heights, so traps at one height only are not suitable for comparing the relative abundance of different species. There was less variation in numbers caught on traps at 3–4 ft. above the ground than on traps at 5–6 ft. or at ground-level.
During the first fortnight of July an average of 200 aphides, 1.6% of which were Myzus persicae , was caught on traps in the position of potato plants. Most of these were not potato aphides, but it is suggested that some of them might act as vectors of potato viruses. The number of aphides caught was roughly proportional to the volume of free air space around the traps, and as the foliage increased in density the numbers caught on lower traps decreased in proportion.
Wind-speed records within and above the crop are discussed and it is shown that conditions are often suitable for voluntary aphis flight in and near the crop on relatively windy days. Records of catches on the half-traps facing the wind and on the other halves showed that voluntary flight decreased with increasing height.
Brilliant yellow traps, coated with adhesive grease, caught more aphides than white traps, which in turn caught more than black.     

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.