Fastest Flyer

University of Florida Book of Insect Records
Chapter 1 Fastest Flyer

T. J. Dean

School of Physics, University of New South Wales at the Australian Defence Force Academy

1 April 2003

The insects with the highest reliably measured airspeeds are desert locusts (Schistocerca gregaria) and corn earworm moths (Helicoverpa zea). These fly at average airspeeds of 33 and 28 km/h respectively (about 21 and 17 mph). Many insects surely fly faster, but their airspeeds have yet to be studied with modern methods. The highest sustained ground speed recorded is that of the black cutworm (Agrotis ipsilon) which flies at speeds of between 97 and 113 km/h (60-70 mph) (Showers & Sappington 1992). Insect airspeed is affected by mass, size, age, gender, feeding, water content, activity type, temperature, humidity, solar radiation, wind, oxygen level, ascent angle and even habitat isolation.

The speed attainable by insects is currently poorly understood (Gauthreaux et al. 1998); indeed Dudley (1997) states that insect airspeed is one of the least known features of flight performance. This paper details the current state of insect flight speed measurements and includes the most complete table of measured speeds to date.

Methods

Stevenson et al.(1995) attribute the large range of flight speeds measured being due to the different methodologies used (e.g. timing with stopwatches; wind tunnels; flight mills etc.). Insects flying freely often have higher speeds than those that are confined to small cages or tethered (although Wagner (1986) states that cage size has no influence on basic flight performance). While measurements have been made for insects flying up pheromone plumes, they are generally slower than freely flying insects (Kuenen & Carde 1993) and their speed decreases with proximity (Willis et al. 1991) and strength of the source (e.g. Meats & Osborne 2000; March & McNeil 2000).

Riley et al. (1997) found that insects flying on flight mills partitioned their effort between lift and thrust substantially differently from those in free flight. Cooter & Armes (1997), Gatehouse & Hacket (1980), and Gatehouse & Woodrow (1987)) consider that mills do not adversely affect the behaviour of the cotton bollworm (Helicoverpa armigera). They do emphasize, however, that the results are meaningful only when used to provide comparative estimates of flight performance between experimental treatments. Other methods of monitoring insect activity include cameras (e.g. El-Sayed et al. 2000; Fry et al. 2000; Noldus et al. 2002; Hardie & Powell 2002), Doppler-radar autocorrelation analysis (e.g. Buchan & Satelle 1979; Sohal & Buchan 1981; Buchan & Moreton 1981; Renou et al. 1999; Knoppien et al. 2000), telescopic observation against the moon (e.g. Preuss & Preuss 1971), and roundabouts (e.g. Michel et al. 1977). For a full review of remote-sensing, telemetric and computer-based technologies see Reynolds & Riley (2002).

In addition to effects of the methods employed on the measured speed, speed has been found to vary between laboratory and naturally reared specimens. McKibben et al. (1988) found that naturally reared cotton boll weevils (Anthonomus grandis) specimens had an average speed 1.2 times greater than that of the laboratory reared specimens, although this was not found for six generations of the fruit fly Ceratitis capitata (Economopoulos 1992). Significant variation has also been found within single species; a total of seven authors have made over 400 measurements of the speed of the desert locust (Schistocerca gregaria) and found ground speeds ranging from 3 to 33 km/h (Table 1).

Results

The most famous and oft-quoted insect flight speed is that of the deer botfly, reputed to be able to fly at over 1,287 km/h (Townsend 1926). However, Langmuir (1938) refuted this claim and calculated that to attain this speed (equivalent to half a horse power) the fly would have to consume 1.5 its own weight in fuel every second. Further observations by Langmuir (1938) found that the maximum speed was more likely to be around 40 km/h. Some of the difficulties with these early measurements arose form the difficulty in separating airspeed from ground speed. Airspeed is the speed relative to the air whereas the ground speed (often the quantity actually measured) is the speed relative to the ground (Fig. 1). This difficulty in separating airspeeds from ground speeds makes some early measurements obtained using ‘less than conventional’ methods (e.g., comparisons with the speed of trains [Twinn et al. 1948]) useless for comparison purposes. This is why the reported speed of 98 km/h for Austrophlebia costalis (Hocking 1953) is not included here.

Figure 1: A vector diagram showing the relationship between insect airspeed, wind speed and ground speed.

The highest airspeeds reported in refereed literature obtained using a reliable method are those of the desert locust (Schistocerca gregaria), 15 individuals having an average speed of reached 33 ± 3 km/h (mean ± SE) (Waloff 1972), and the corn earworm moth (Helicoverpa zea), ten individuals reaching an average speed of 28 ± 8 km/h (Quero et al. 2001). In the unrefereed literature, a noteworthy record is that of a male horsefly (Hybomitra hinei) that was estimated to achieve an airspeed of around 145 km/h (89 mph) while chasing an air rifle pellet (Kunzig 2000).

The highest sustained ground speed recorded is that of the black cutworm (Agrotis ipsilon) which flies at speeds of between 97 and 113 km/h at heights of between 300 and 600 m ‘riding’ on winds ahead of cold fronts (Showers & Sappington 1992) although this species only has an airspeed of between 9 and 13 km/h (Jia & Cao 1992).

Discussion

Insect flight speed has been found to be affected by the followin insect characteristics:

1. mass (e.g. Dudley & Srygley 1994; Dudley 1997),
2. size (e.g. Larkin 1991; Fischer & Kutsch 2000),
3. age (e.g. Karlsson 1994; Banjaree 1988),
4. gender (e.g. Rogowitz & Chappell 2000; Willmott & Ellington 1997; Lingren et al. 1995),
5. amount of feeding (e.g. David 1978; Fadamiro & Wyatt 1995),
6. water content (e.g. Danks 2000; Lehmann et al. 2000),
7. activity type (e.g. David & Hardie 1988; May 1999).

Also, insect flight speed is affected by the following environmental factors:

1. temperature (e.g. Gilchrist et al. 1997; Isard et al. 2000; Fitzgerald & Underwood 2000; Elliott et al. 2000),
2. humidity (e.g. Gunn 1937; Pielpou & Gunn 1940; Dorner & Mulla 1962),
3. solar radiation (e.g. Rudinsky & Vite 1956; Ostrand et al. 2000; Carde & Knowls 2000; Vicens & Bosch 2000; Lloyd 2000; Schneider 1965),
4. wind (e.g. Aluja et al. 1993; Hardie & Young 1997),
5. oxygen levels (e.g. Ellington et al. 1990; Joos et al. 1997; Harrison & Lighton 1998; Dekker et al. 2001),
6. habitat isolation (e.g. Denno et al. 2001), and
7. ascent angle (Kutsch et al. 1999).

Given the wide variety of possible effects on insect flight speed, studies of large numbers of insects from a single species have found that the distribution of speeds approximately follows a normal distribution (e.g. Tuxhorn & McShaffrey 1998; Nachtigall 2001; Dean & Drake 2002). Most insects have airspeeds of less than 21 km/h. All currently available measured insect speeds are listed in Table 1. It should be borne in mind, however, that many of the early measurements were made using inaccurate methods and may reflect only a single speed measurement.

References

Abbott, C.H., 1951, A quantitative study of the migration of the painted lady butterfly, Vanessa cardui L.: Ecology, 32, 155-171.

Aluja, M., Prokopy, R.J., Buonaccorsi, J.P., and Carde, R.T., 1993, Wind tunnel assays of olfactory responses of female Rhagoletis pomonella flies to apple volatiles; effect of wind speed and odour release rate: Entomologia Experimentalis et Applicata, 68, 99-108.

Baker, P.S., and Cooter, R.J., 1979, The natural flight of the migratory locust, Locusta migratoria L.: J. Comp. Physiol., 131, 79-87.

Baker, P.S., Gewecke, M., and Cooter, R.J., 1981, The natural flight of the migratory locuts, Locusta migratoria L.III. Wing-beat frequency, flight speed and attitude: J. comp. Physiol., 141, 233-237.

Baker, T.C., Willis, M.A., and Phelan, P.L., 1984, Optomotor anemotais polarizes self-steered zigzagging in flying moths: Physiological Entomology, 9, 365-376.

Barata, E.N., and Araujo, 2001, Olfactory, orientation responses of the eucalyptus woodborer, Phoracantha semipunctata, to host plant in a wind tunnel: Physiological Entomology, 26, 26-37.

Balciunas, J., and Knoff, K., 1977, Orientation, flight speeds, and tracks of three species of migrating butterflies: The Florida Entomologist, 60, 37-39.

Banjeree, S., 1988, Organisation of wing cuticle in Locusta migratoria Linnaeus, Tropidacris cristata Linnaeus and Romalea microptera Beauvais: International Journal of Insect Morphology & Embryology, 17, 313-326.

Bentley, C.A., 1914, Notes on experiments to determine the reactions of mosquitos to artificial light: indian J. med. res. Suppl., 5, 9-11. (Cited in Hocking, 1953).

Berrigan, D., and Lighton, J.R.b, Bioenergetic and kinematic consequences of limblessness in larval diptera: J. exp. Biol., 179, 245-259.

Betts, C.R., and Wootton, R.J., 1988, Wing shape and flight behaviour in butterflies (Lepidoptera: Papilionoidea and Hesperioidea): a preliminary analysis: J. exp. Biol., 138, 271-288.

Beutler, R., 1950, Zeit und raum in leben der sammelbiene: Naturwissenschaften, 37, 102-105.

Bodenheimer, F.S., 1931, Ueber die Temperaturabhangigkeit von Isekten: III. Die beziehungen der vorzugstemperatur zur luftfeuchtigkeit der umgebung: Zeitschr. vergl. Physiol., 13, 740-747 (cited in Dorner and Mulla, 1962).

Brackenbury, J., 1999, Water skating in the larvae of Dixella aestivalis (Diptera) and Hydrobius fuscipes (coleoptera): J. exp. Biol., 202, 845-853.

Brady, J., 1991, Flying mate detection and chasing by tsetse flies (Glossina): Physiological Entomology, 16, 153-161.

Briegel, H., Knusel, I., and Timmermann, S.E., 2001, Aedes aegypti: size, reserves, survival and flight performance: Journal of Vector Ecology, 26, 21-31.

Buchan, P.B., and Moreton, R.B., 1981, Flying and walking of small insects (Mustica domestica) recorded differentially with a standing-wave radar actograph: Physiological Entomology, 6, 149-155.

Buchan, P.B., and Sattelle, D.B., 1979, A radar-doppler autocorrleation analysis of insect activity: Physiological Entomology, 4, 103-109.

Byers, J.A., 1996, An encounter rate model of bark beetle populations searching at random for susceptible host trees: Ecological Modelling, 91, 57-66.

Byrne, D.N., 1999, Migration and dispersal by the sweet potato whitefly, Bemisia tabaci: Agricultural and Forest Meteorology, 97, 309-316.

Callahan, P.S., 1965, A photoelectric-photographic analysis of flight behaviour in the Corn Earworm, Heliothis zea and other moths: Annals of the Entomological Society of America, 58, 159-169.

Capaldi, E.A., Smith, A.D., Osborne, J.L., Fahrbach, S. E., Farris, S. M., Reynolds, D. R., Edwards, A. S., Martin, A., Robinson, G.E, Poppy, G.M and Riley, J.R., 2000, Ontogeny of orientation flight in the honeybee revealed by harmonic radar: Nature, 403, 537-540.

Carde, R.T., and Knols, G.J., 2000, Effects of light levels and plume structure on the orientation manoeuvres of male gypsy moths flying along pheremone plumes: Physiological Entomology, 25, 141-150.

Chaudhari, G.B., Bharpdoa, T.M., Patel, J.J>, Patel, K.I., and Patel, J.R., 1999, Effect of weather on activity of cotton bollworms I middle Gujarat: Journal of Agrometeorology, 1, 137-142.

Chen, R.C., Wu, J.R., Zhu, S.D., and Zhang, J.X., 1984, Flight capacity of the brown planthopper Nilaparvata lugens Stal.: Acta Entomologica Sinica, 27, 121-127.

Chiba, Y., Uki., M., Kawasaki, Y., Matsumoto, A., and Tomioka, K., 1993, Entrainability of circadian activity of the mosquito Culex pipiens pallens to 24-hr temperature cycles, with special reference to involvement of multiple oscillators: J. Biol. Rhythms, 8, 211-220.

Cloudsley-Thompson, J.L., 1989, Temperature and the activity of ants and other insects in central Australia: Journal of Arid Environments, 16, 185-192.

Coelho, J.R., and Holliday, C.W., 2001, Effects of size and flight performance on intermale mate competition I the cicada killer: Sphecius speciosus Drury (Hymenoptera: Sphecidae): Journal of Insect Behaviour, 14, 345-351.

Collett, T.S., and Land. M.F., 1975, Visual control of flight behaviour in the hoverfly, Syritta pipiens L.: J. comp. Physiol., 99, 1-66.

Colvin, J., Brady, J., and Dransfield, R.D., 1989, Visually-guided, upwind turning behaviour of free-flying tsetse flies in odour-laden wind: a wind-tunnel study: Physiological Entomology, 14, 31-39.

Cooter, R.J., and Armes, N.J., 1993, Tethered flight technique for monitoring the flight perforamcne of Helicoverpa armigera (Lepidoptera: Noctuidae) : Population Ecology, 22, 339-345.

Correale, S., and Crocker, R.L., 1976, Ground speed of 3 species of migrating lepidoptera: The Florida Entomologist, 59, 424.

Danks, H.V., 2000, Dehydration in dormatn insects: Journal of Insect Physiology 46, 837-852.

David, C.T., 1978, The relationship between body angle and flight speed in free-flying Drosophila: Physiological entomology, 3, 191-195.

David, C.T., 1982, Compensation for height in te control of groundspeed by Drosophila in a new, ‘barber’s pole’ wind tunnel: J. Comp. Physiol., 147, 485-193.

David, C.T., and Hardie, J., 1988, The visual responses of free-flying summer and autumn form of the black bean aphid, Aphis fabae, in an automated flight chamber: Physiological Entomology, 13, 277-284.

Dean, T.J., and Drake, V.A., 2002, Properties of biotic tragets observed with an X-band radar profiler and the potential for bias in winds retrieved from Doppler weather radars: Proceedings of the 11th Australasian Remote Sensing and Photogrammetry Conference, 698-711.

Dekker, T., Takken, W., and Carde, R.T., 2001, Structure of host-odour plumes influences catch of Anopheles gambiae s.s. and Aedes aegypti in a dual-choice olfactometer: Physiological Entomology, 26, 124-134.

Demoll, R., 1918, Der Flug der Insekten und der Vogel: Jena. (Cited in Hocking, 1953).

Denno, R.F., Hawthorne, D.J., Throne, B.L., and Gratton, C., 2001, Reduced flight capability in British Virgin Island populations of a wing-dimorphic insect: the role of habitat isolation, persistence and structure: Ecological Entomology, 25, 25-26.

DeVries, P.J., and Dudley, R., 1990, Morphometrics, airspeed, thermoregulation, and lipid reserves of migrating Urania fulgens (Uraniidae) moths in natural free flight: Physiological Zoology, 63, 235-251.

Dorner, R.W., and Mulla, S., 1962, Laboratory study of wind velocity and temperatur preference of Hippelates eye gnats: Annals of the Entomological Society of America, 55, 36-39.

Duan, J.J., Weber, D.C., Hirs, B., and Corn, S., 1996, Spring behavioural patterns of the apple blossom weevil: Entomolgia Experimentalis et Applicata, 79, 9-17.

Dudley. R., 1997, The biomechanics of insect flight; form function, evolution: Princeton University Press.

Dudley, R. and Srygley, R.B., 1994, Flight physiology of neotropical butterflies: Allometry of airspeeds during natural free flight: The Journal of Experimental Biology, 191, 125-139.

Economopoulis, A.P., 1992, Adaptation of the Mediterranean fruit fly (Diptera: Tephritidae) to artificial rearing: Journal of Economic Entomology, 85, 753-758.

El-Sayed, A., Godde, J., and Arn, H., 2000, A computer-controlled video system fro real-time recording of insect flight int hree dimensions: Journal f Insect Behaviour, 13, 881-900.

Ellington, C.P., 1991, Limitations on animal flight performance: J. exp. Biol., 160, 71-91.

Ellington, C.P., Machin, K.E., and Casey, T.M., 1990, Oxygen consumption of bumblebees in forward flight: Nature, 347, 472-473.

Elliott, N.C., Kieckhefer, R.W., and Beck, D.A., 2000, Adult Coccinellid activity and predation on Aphids in Spring cereals: Biological control, 17, 218-226.

Fadamiro, H.Y., 1996, Flight and landing behaviour of Prostephanus truncatus (Horn) (Coleoptera: Bostrichidae) in relation to wind speed: Journal of Stored Products Research, 32, 233-238.

Fadamiro, H.Y., 1996b, Influence of stimulus dose and wind speed on the orientation behaviour of Prostephanus truncatus (Coleoptera: Bostrichidae) to pheromone: Bulletin of Entomological Research, 86, 659-665.

Fadamiro, H.Y., 1997, Free flight capacity determination in a sustained flight tunnel: Effects of age and sexual state on the flight duration of Prostephanus truncatus: Physiological Entomology, 22, 29-36.

Fadamiro, H.Y., and Wyatt, T.D., 1995, Flight inititation by Prostephanus truncatus in relation to time of day, temperature , relative humidity and starvation: Entomologia Experimentalis et Applicata, 75, 273-277.

Fadamiro, H.Y., Wyatt, T.D., and Birch, M.C., 1998, Flying beetles respond to moths predict: optomotor anemotaxis to pheromone plumes at different heights: Journal of Insect Behaviour, 11, 549-557.

Fischer, H., and Ebert, E., 1999, Tegula function during free locust flight in relation to motor pattern, flight speed and aerodynamic output: J. Exp. Biol., 202, 711-721.

Fischer, H., and Kutsch, W., 1999, Timing of elevator muscle activiy during climbing in free locust flight: J. exp. Biol., 202, 3575-3586.

Fischer, H., and Kutsch, W., 2000, Relationships between body mass, motor output and flight variables during free flight of juvenile and mature adult locusts, Schistocerca gregaria: J. exp. Biol., 203, 2723-2735.

Fitzgerald, T.D.,a nd Underwood, D.L.A., 2000, Winter foraging patterns and voluntary hypothermia int he social caterpillar Eucheira socialis: Ecological Entomology 25, 35-44.

Frisch, K. von., 1927, Aus dem Leben der Bienen: Berlin. (Cited in Hocking, 1953).

Fry, S.N., Bischel, M., Mueller, P., and Robert, D., 2000, Tracking of flying insects using pan-tilt cameras: Journal of Neuroscience Methods, 101, 59-67.

Gatehouse, A.G., and Hackett, D.S., 1980, A technique for studying lfight behaviour of tethered Spodoptera exempta moths: Physiological Entomology, 5, 215-222.

Gatehouse, A.G., and Woodrow, K.P., 1987, Simultaneous monitoring of flight and oviposition of individual velvetbean caterpillar moths (by Wales, Barfiels and Leppla, 1985); a critique : Physiological Entomology, 12, 117-121.

Gauthreaux, S. A. Jr., Mizrahe, D.S., and Belser, C. G., 1998b, Bird migration and bias of WSR-88D wind estimates: Weather and Forecasting, 13, 465-481.

Gewecke, ?, Control of flying speed in locusts and its significance for their migrations:.