Chapter 1: Fastest Flyer

Read Editor's Note

T.J. Dean
School of Physics, University of New South Wales at the Australian Defence Force Academy
April 1, 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 calculating 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 the maximum speed was more likely to be around 40 km/h. Some of the difficulties with early measurements arose from the difficulty separating airspeed from ground speed. Airspeed is the speed relative to the air whereas ground speed (often the quantity actually measured) is the speed relative to the ground (Figure 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: 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 following 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; Quero et al. 2001; 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; Pielou & 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)
  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.

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Table 1

Published insect speeds in ms-1, errors in brackets (1 m/s = 3.6 km/h = ca. 2.2 mph).

No.: number

Range:
 A: average, X: max, N: min, R: range

Type:
 A: air-speed, G: ground-speed.
 This was difficult to determine for some (especially early) studies and readers are advised
 to check the original reference.

Method:
 WT: wind tunnel; FF: free flying; FC: flight chamber; T: tethered; M: Mill.

Coleoptera

Family Genus species Speed No. / Range / Type / Method Source
Bostrichidae Prostephanus truncatus 0.2   / A / A / WT Fadamiro, 1996
Prostephanus truncatus (20 m/s wind) 0.07 (0.01) 15 / A / A / WT Fadamiro, 1996b
Prostephanus truncatus (25 m/s wind) 0.08 (0.01) 15 / A / A / WT Fadamiro, 1996b
Prostephanus truncatus (32 m/s wind) 0.1 (0.01) 9 / A / A / WT Fadamiro, 1996b
Ceramycidae Phoracantha semipunctata (in plume) 0.66 (0.21) 32 / A / A / WT Barata & Araujo, 2001
Phoracantha semipunctata (outside plume) 0.52 (0.14) 40 / A / A / WT Barata & Araujo, 2001
Cicindelidae Cicindera hybrida 0.6   / A / A / FF Nachtigall, 1996
3   / X / A / FF Nachtigall, 1996b
Dytiscidae unknown 1.97   / ? / G / ? Lane, 1941
Lampyridae Photinus collustrans 1.2 123 / A / G / FF Lloyd, 2000
0.14-2.1 123 / A / G / FF Lloyd, 2000
Nitidulidae Carpophilus hemipterus (f) 0.44   / A / A / T Wu & Laughlin, 1994
Carpophilus hemipterus (m) 0.33   / A / A / T Wu & Laughlin, 1994
Scolytidae Dendroctonus pseudotsugae 0.34-0.72   / A / A / T Rudinsky & Vite, 1956
Ips typographus 0.15   / N / G / FF Zolubas & Skirkevicius, 1992
various 2   / X / G / FF Byers, 1996

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Diptera

Family Genus species Speed No. / Range / Type / Method Source
Calliphoridae Calliphora vicina 1.2   / X / A / FC Schilstra & Hateren, 1999a;b
1   / X / A / FC Schilstra, 1999
1.2   / X / A / FC Schilstra, 1999
Phaenicia sericata 1.45-2.05   / A / A / FC Yurkiewicz & Smyth, 1966
Cecidomyidae Mayetiola destructor 7   / A / G / FC Lewis & Taylor, 1967
Chloropidae Hister merdarius 1.92   / A / G / FC Lewis & Taylor, 1967
Oscinella frit 0.76   / A / G / FC Lewis & Taylor, 1967
Culicidae Aedes aegypti (female) 0.18   / A / A / M Briegel et al., 2001
Aedes aegypti (poor) 0.13   / A / A / M Briegel et al., 2001
Aedes aegypti (fast) 0.28   / A / A / M Briegel et al., 2001
0.33   / X / A / M Briegel et al., 2001
Aedes aegypti 1.5   / A / G / FF Kennedy, 1939
0.08-0.33   / A / A / FF Kennedy, 1939
Aedes flavescens 1.1 3 / A / A / M Hocking, 1953
Aedes nearcticus 1.0 10 / A / A / M Hocking, 1953
Anopheles fuliginosus 0.08-0.33   / A / A / FC Bentley, 1914
Anopheles melas 1.2   / X / G / FF Snow, 1980
Culex tarsalis 2.15   / A / G / FF Bailey et al., 1965
Culex thalassius 1.2   / A / G / FF Snow, 1980
unknown 1.39   / ? / G / ? Lane, 1941
Drosophilidae Drosophila hydei (fed) 0.3 (0.085)   / A / G / WT David, 1978
Drosophila hydei (starved) 0.46 (0.11)   / A / G / WT David, 1978
0.5   / A / A Lehmann, 2001
Drosophila virilis 2   / A / A / WT Vogel, 1967
2   / A / A / T Vogel, 1966
Drosophila melangaster 1.37   / A / G / FC Lewis & Taylor, 1967
0.70 (0.01) 113 / A / A / WT Marden et al., 1997
0.64 (0.02) 88 / A / A / WT Marden et al., 1997
0.46 (0.02) 99 / A / A / WT Marden et al., 1999
0.52 (0.01) 144 / A / A / WT Marden et al., 2000
Drosophila 1.4 26 / A / A / M Hocking, 1953
Empididae Rhamphomyia 3.05   / A / G / FC Lewis & Taylor, 1967
Equidae Hippelates collusor 0.9   / A / A / WT Dorner & Mulla, 1962
Fanniidae Fannia canicularis 0.4-0.8   / A / A / FC Nachtigall, 1999
0.5-0.65   / A / A / FC Nachtigall, 1999b
Glossinidae Glossina morsitans morsitans 1.8   / A / G / WT Colvin et al., 1989
4.8 (0.1)   / A / G / FF Brady, 1991
Muscidae Musca domestica 2.23   / ? / G / ? Lane, 1941
3.05   / A / G / FC Lewis & Taylor, 1967
1.8   / A / G / FF McKeown, 1944
Stomoxys calcitrans 0.33   / A / A / FF Bailey et al., 1973
0.55   / A / A / FF Eddy et al., 1962
Musca sp. 0.28 (0.032) 8 / A / G / FF Golding et al., 2001
Oestridae Tracheomyia macropi 11.17   / X / G / ? Lane, 1941
Psychodidae Phlebotomus ariasi 0.65-0.70   / X / A / WT Killick-Kendrick et al., 1986
Schizophora Calliphoridae 2.68   / ? / G / ? Lane, 1941
Simuliidae Simulium venustum 1.5 16 / A / A / M Hocking, 1953
2.4 3 / A / A / M Hocking, 1953
Syrphidae Eristalis tenax 0.19 (0.03) 8 / A / G / FF Golding et al., 2001
Syritta pipiens 1.4-4.6   / A / A / FC Collet & Land, 1975
Syrphus ribesii 0.19 (0.01) 9 / A / G / FF Golding et al., 2001
Tabanidae Hybomitra hinei 40.3   / X / G / FF Kunzig, 2000
Tabanids (unspecified) 18   / N / G / FF Schwardt, 1932
Tabanus affinis 2.3 71 / A / A / T&M Hocking, 1953
6.5   / A / G / FC Lewis & Taylor, 1967
12.5   / X / G / FF Twinn et al., 1948
Tabanus bovinus 14   / X / A / FF Demoll, 1918
4   / A / A / T Magnan, 1934
Tabanus septentrionalis 2.4 10 / A / A / M Hocking, 1953
6.5   / A / G / FC Lewis & Taylor, 1967
various 13.4   / ? / G / ? Lane, 1941
various 12.5   / A / G / FF McKeown, 1944

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Ephemeroptera

Family Genus species Speed No. / Range / Type / Method Source
unknown unknown 0.5   / ? / G / ? Lane, 1941

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Hemiptera

Family Genus species Speed No. / Range / Type / Method Source
Aphididae Aphis fabae 0.76   / A / G / FC Lewis & Taylor, 1967
0.13-0.28   / A / A / WT Kennedy & Thomas, 1974
various 0.54-0.67   / A / A / WT Haine, 1955
various 3.15 (0.19)   / A / A / WT Hardie & Young, 1997
various 2.76 (0.19)   / A / A / WT Hardie & Young, 1997
Cicadellidae Circulifer tenellus 0.27-0.85   / A / A / WT Lawson et al., 1951
Miridae Lygus lineolaris 0.45   / A / A / M Stewart & Gaylor, 1994
various various 3   / A / G / FF Nilakhe & Buainain, 1988

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Homoptera

Family Genus species Speed No. / Range / Type / Method Source
Aleyrodidae Aleyrodes 0.46   / A / G / FC Lewis & Taylor, 1967
Bemisia tabaci 0.04-0.16   / A /   / FC Byrne, 1999
0.16-0.23 30 / A / G / WT Isaacs et al., 1999
0.16-0.49 30 / A / A / WT Isaacs et al., 1999
Aphididae Drepanosiphum platanoidis 0.91   / A / G / FC Lewis & Taylor, 1967
Jacksonia papillata 0.37   / A / G / FC Lewis & Taylor, 1967
Macrosiphum pisum 0.55   / A / G / FC Lewis & Taylor, 1967
Cicadellidae Typhlocyba 1.16   / A / G / FC Lewis & Taylor, 1967
Delphacidae Nilaparvata lugens 22.4   / X / A / M Chen et al., 1984

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Hymenoptera

Family Genus species Speed No. / Range / Type / Method Source
Apiidae unknown 3-15.7 100 / A / G / FF Osborne et al., 1999
unknown 4.9   / ? / G / ? Lane, 1941
Apis mellifera 2.6 16 / A / A / M Hocking, 1953
4.14   / A / G / FC Lewis & Taylor, 1967
3.3-5.1   / A / A / T Nachtigall et al., 1995
0.19 (0.025) 11 / A / G / FF Golding et al., 2001
2.2-3.6   / A / G / FF McKeown, 1944
Apis mellifera (drone) 1.8   / A / G / FF Park, 1923
1.8   / A / A / FF Schnell-larsen, 1934
Apis mellifera (Worker - empty) 8.3   / A / A / T Hansson, 1945
4.9   / ? / G / ? Lane, 1941
5.6   / A / G / HR Capaldi et al., 2000
3.6   / A / G / HR Capaldi et al., 2000
Apis mellifera (Worker - full) 7.2   / A / A / T Hansson, 1945
2.4   / ? / G / ? Lane, 1941
Apis mellifera (Worker) 5.56   / A / G / FF Beutler, 1950
3.75   / A / G / FF Demoll, 1918
2.5   / A / A / T Magnan, 1934
6.12   / A / G / FF Park, 1923
6   / A / A / T von Frisch, 1927
14   / A / G / FF Wenner, 1963
Ichneumonidae Ichneumon 1.67   / A / G / FC Lewis & Taylor, 1967
Vespidae Vespa crabro 5.9   / ? / G / ? Lane, 1941
unknown unknown 5.4   / ? / G / ? Lane, 1941

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Lepidoptera

Family Genus species Speed No. / Range / Type / Method Source
Hesperiidae Chioides catillus 3.6 (0.2) 2 / A / A / FF Srygley, 2001
Dryas julia 3.1 (0.6) 6 / A / A / FF Srygley, 2001
unknown 5.1 1 / A / A / FF Srygley, 2001
Urbanus simplicius 3.5 1 / A / A / FF Srygley, 2001
unknown 5.3 1 / A / A / FF Srygley, 2001
Calpodes ethlius 7.5 2 / A / A / FF Dudley & Srygley, 1994
Cyclosemia anastomosis 5.7 1 / A / A / FF Dudley & Srygley, 1994
Heliopetes sp. 4.1 2 / A / A / FF Dudley & Srygley, 1994
Typhedanus undulatus 7.4 1 / A / A / FF Dudley & Srygley, 1994
Urbanus proteus 4.5 98 / A / G / FF Balciunas & Knopf, 1977
6.1 98 / A / A / FF Balciunas & Knopf, 1977
Libytheidae Libythia bachmani 4.47-6.7   / A / G / FF Parman, 1926
Lycaenidae Lycaena phlaeas 2.86   / A / G / FC Lewis & Taylor, 1967
Lymantriidae Lymantria dispar 4.5 24 / A / G / WT Kuenen & Carde, 1993
4.7 24 / A / G / WT Kuenen & Carde, 1993
5.3 24 / A / G / WT Kuenen & Carde, 1993
0.64 (0.18) 20 / A / A / WT Carde & Knols, 2000
0.53 (0.06) 20 / A / A / WT Carde & Knols, 2000
0.48 (0.06) 20 / A / A / WT Carde & Knols, 2000
Noctuidae Agrochola lychnidis 4.35   / A / G / FC Lewis & Taylor, 1967
Agrotis ipsilon 26.8-31.3   / A / G / T Showers & Sappington, 1992
Schistocerca gregaria 3.5 47 / A / G / FF Lingren et al., 1995
7.9 (2.3) 10 / A / A / WT Quero et al., 2001
Schistocerca gregaria (casting) 5.7 (2.4) 10 / A / A / WT Quero et al., 2001
Noctuidae & Sphingidae unknown 4.47-6.7   / A / A / FC Callahan, 1965
Nymphalidae Adelpha iphiclus 3.6 1 / A / A / FF Dudley & Srygley, 1994
Agranlis vanillae 3.6 1 / A / A / FF Srygley, 2001
2.8 2 / A / G / FF Correale & Crocker, 1976
5 1 / A / A / FF Dudley & Srygley, 1994
Anartia fatima 3.6 7 / A / A / FF Dudley & Srygley, 1994
Anartia fatima (females) 2.6 (0.2) 5 / A / A / FF Srygley & Kingsolver, 2000
Anartia fatima (males) 2.5 (0.4) 27 / A / A / FF Srygley & Kingsolver, 2000
Archaeoprepona demophon 7 1 / A / A / FF Dudley & Srygley, 1994
Cissia sp. 2.2 1 / A / A / FF Dudley & Srygley, 1994
Danaus eresimus 3.9 2 / A / A / FF Dudley & Srygley, 1994
3 2 / A / A / FF Srygley, 2001
Danaus gilippus 3.6 4 / A / A / FF Dudley & Srygley, 1994
Danaus plexippus 4.9   / A / G / FF Urquhart, 1960
8.9-11.2   / A / G / FF Urquhart, 1960
Danaus plexippus (parasite infected) 0.9622 20 / A / A / M Yueh, 1999
0.9345 20 / A / A / M Yueh, 1999
Dione juno 3.8 3 / A / A / FF Dudley & Srygley, 1994
Dryadula phaetusa 3.7 8 / A / A / FF Dudley & Srygley, 1994
Dryas iulia 4.5 5 / A / A / FF Dudley & Srygley, 1994
Eueides lybia 2.3 1 / A / A / FF Dudley & Srygley, 1994
Eunica alcmena 3 1 / A / A / FF Dudley & Srygley, 1994
Eunica sp. 4.8 1 / A / A / FF Dudley & Srygley, 1994
Hamadryas feronia 4.4 3 / A / A / FF Dudley & Srygley, 1994
Hamadryas ipthime 4.5 1 / A / A / FF Dudley & Srygley, 1994
Heliconius charitonius 2.3 1 / A / A / FF Dudley & Srygley, 1994
Heliconius eleuchia 1.9 1 / A / A / FF Dudley & Srygley, 1994
3.3 1 / A / A / FF Dudley & Srygley, 1994
Heliconius erato 1.7 3 / A / A / FF Dudley & Srygley, 1994
Heliconius hecale 2.1 7 / A / A / FF Dudley & Srygley, 1994
Heliconius melpomene 2.6 1 / A / A / FF Dudley & Srygley, 1994
Heliconius sapho 2 1 / A / A / FF Dudley & Srygley, 1994
Heliconius sara 2.7 7 / A / A / FF Dudley & Srygley, 1994
Historis acheronta 6.5 11 / A / A / FF Dudley & Srygley, 1994
4.3 1 / A / A / FF Srygley, 2001
Junonia evarete 2.3 2 / A / A / FF Dudley & Srygley, 1994
3.8 1 / A / A / FF Srygley, 2001
Laparus doris 2.7 1 / A / A / FF Dudley & Srygley, 1994
Marpesia chiron 4.6 23 / A / A / FF Dudley & Srygley, 1994
Marpesia petreus 4.9 13 / A / A / FF Dudley & Srygley, 1994
Mechanitis lysimnia 0.7 1 / A / A / FF Dudley & Srygley, 1994
Mechanitis polymnia 2.5 1 / A / A / FF Dudley & Srygley, 1994
Melinaea parallelis 2.5 2 / A / A / FF Dudley & Srygley, 1994
Melinaea scylax 3.5 1 / A / A / FF Dudley & Srygley, 1994
Morpho amathonte 2.6 1 / A / A / FF Dudley & Srygley, 1994
Olyras insignis 2.5 1 / A / A / FF Dudley & Srygley, 1994
Philaethria dido 2.5 5 / A / A / FF Dudley & Srygley, 1994
Precis coenia 2.8 10 / A / G / FF Balciunas & Knopf, 1977
5.3 10 / A / A / FF Balciunas & Knopf, 1977
Precis lavinae 2.7 7 / A / G / FF Correale & Crocker, 1976
Pyrrhogyra crameir 2.2 1 / A / A / FF Dudley & Srygley, 1994
Pyrrhogyra neaerea 1.8 1 / A / A / FF Dudley & Srygley, 1994
Pyrrhogyra otolais 3.6 1 / A / A / FF Dudley & Srygley, 1994
Siproeta stelenes 3.7 2 / A / A / FF Dudley & Srygley, 1994
Vanessa atalanta 3.9   / ? / G / ? Lane, 1941
Vanessa cardui 2.2-6.1   / A / G / FF Abbott, 1951
4.9-8   / A / G / FF Williams, 1930
Papilionidae Battus polydamas 5.8 3 / A / A / FF Dudley & Srygley, 1994
Eurytides dolicaon 4.5 1 / A / A / FF Dudley & Srygley, 1994
Eurytides ilus 4.5 3 / A / A / FF Dudley & Srygley, 1994
Eurytides protesilaus 5.3 3 / A / A / FF Dudley & Srygley, 1994
Papilio anchisiades 4.6 2 / A / A / FF Dudley & Srygley, 1994
Papilio thoas 4.6 6 / A / A / FF Dudley & Srygley, 1994
Parides arcas 2.9 3 / A / A / FF Dudley & Srygley, 1994
Parides sesostris 3.1 2 / A / A / FF Dudley & Srygley, 1994
Pieridae Anteos maerula 5.4 4 / A / A / FF Srygley, 2001
Aphrissa boisduvallii 5.1 29 / A / A / FF Dudley & Srygley, 1994
Aphrissa statira 5.3 36 / A / A / FF Dudley & Srygley, 1994
4 3 / A / A / FF Srygley, 2001
Appias drusilla 4 5 / A / A / FF Dudley & Srygley, 1994
4 1 / A / A / FF Srygley, 2001
Ascia monuste 3.1-4.5   / A / G / FF Nielsen, 1961
3.4 14 / A / A / FF Srygley, 2001
Colia philodice 1.3-2   / A / G / FF Leigh & Smith, 1959
Colias eurytheme 2.45 (0.15) 9 / A / G / FF Srygley & Kingsolver, 1998
Eurema proterpia 2.9 1 / A / A / FF Dudley & Srygley, 1994
Itaballia demophile 2.7 3 / A / A / FF Dudley & Srygley, 1994
Itaballia pandosia 2.2 2 / A / A / FF Dudley & Srygley, 1994
Melete florinda 1.8 1 / A / A / FF Dudley & Srygley, 1994
Phoebis agarithe 4.1 3 / A / A / FF Srygley, 2001
Phoebis argante 4.9 18 / A / A / FF Dudley & Srygley, 1994
Phoebis philea 4.9 6 / A / A / FF Dudley & Srygley, 1994
Phoebis sennae 3.61 66 / A / G / FF Correale & Crocker, 1976
4.1 45 / A / G / FF Balciunas & Knopf, 1977
5.1 45 / A / A / FF Balciunas & Knopf, 1977
6.2 3 / A / A / FF Dudley & Srygley, 1994
4 25 / A / A / FF Srygley, 2001
Phoebis trite 4.8 2 / A / A / FF Dudley & Srygley, 1994
Pieris rapae 2.35 (0.28) 9 / A / G / FF Srygley & Kingsolver, 1998
Pontia occidentalis 2.1 (0.43) 9 / A / G / FF Srygley & Kingsolver, 1998
Sphingidae Manduca sexta 5   / X / A / FC Willmott & Ellington 1997/1997b
Manduca sexta (towards light) 0.4 (0.03) 61 / A / A / FC Spencer et al., 1997
Manduca sexta (away from light) 0.29 (0.02) 61 / A / A / FC Spencer et al., 1997
0.57 38 / A / A / FC Stevenson et al., 1995
1.02 37 / A / A / FC Stevenson et al., 1995
1.58 62 / A / A / FC Stevenson et al., 1995
3.38 93 / A / A / FC Stevenson et al., 1995
22.3   / ? / G / ? Lane, 1941
13.4   / X / G / FF McKeown, 1944
Tortricidae Grapholita molestat 0.80 (0.27)   / A / A / WT Baker et al., 1984
Uraniidae Urania fulgens (female) 3.74 (0.26) 14 / A / A / FF DeVries & Dudley, 1990
Urania fulgens (male) 4.08 (0.21) 13 / A / A / FF DeVries & Dudley, 1990

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Neuroptera

Family Genus species Speed No. / Range / Type / Method Source
Chrysopidae Chrysopa carnea 0.76   / A / G / FC Lewis & Taylor, 1967

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Odonata

Family Genus species Speed No. / Range / Type / Method Source
Aeshnidae Austrophlebia costalis 25-27   / ? / G / ? Lane, 1941
Libellulidae Sympetrum sanguineum 2-5   / A / A / FC Wakeling & Ellington, 1997
Libellula luctuosa 1.86 (0.58) 1805 / A / G / FF Tuxhorn & McShaffrey, 1998
0.41-4.44 1805 / R / G / FF Tuxhorn & McShaffrey, 1998
Libellula lydia 2.12 (0.62) 4411 / A / G / FF Tuxhorn & McShaffrey, 1998
0.68-6.64 4411 / R / G / FF Tuxhorn & McShaffrey, 1998
Pachydiplax longipennis 1.54 (0.55) 59 / A / G / FF Tuxhorn & McShaffrey, 1998
0.53-2.89 59 / R / G / FF Tuxhorn & McShaffrey, 1998
Tramea lacerata 2.61 (0.85) 93 / A / G / FF Tuxhorn & McShaffrey, 1998
1.59-6.88 93 / R / G / FF Tuxhorn & McShaffrey, 1998

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Orthoptera

Family Genus species Speed No. / Range / Type / Method Source
Acrididae Locusta migratora 3.6 (0.2)   / A / A / FC Kutsch & Fuchs, 2000
4.6   / A / G / FF Baker et al., 1981
4.6   / A / G / FF Gewecke
Schistocerca gregaria 4.1 77 / A / A / FC Fischer & Ebert, 1999
4.48 37 / A / A / FF Fischer & Kutsch, 1999
2.5   / N / A / FP Katz & Gosline, 1993
4.5-5   / A / A / T Riley et al., 1988
0.89   / A / A / FF Sayer, 1965
5.36   / A / G / FF Sayer, 1965
3.1 16 / A / A / FF Waloff, 1972
4.3 17 / A / A / FF Waloff, 1972
5.3   / A / A / FF Waloff, 1972
5.0   / A / A / FF Waloff, 1972
5.0 12 / A / A / FF Waloff, 1972
6.1 17 / A / A / FF Waloff, 1972
5.1 11 / A / A / FF Waloff, 1972
8.1 17 / A / A / FF Waloff, 1972
6.1 21 / A / A / FF Waloff, 1972
6.3   / A / A / FF Waloff, 1972
7.8 16 / A / A / FF Waloff, 1972
6.2 23 / A / A / FF Waloff, 1972
9.3 12 / A / A / FF Waloff, 1972
4.9-8   / A / G / FF Waloff & Rainey, 1951
3.5   / A / A / FC Weis-Fogh, 1976
3.0-4.5 158 / A / A / FC Weis-Fogh, 1956
3.5-5.5   / A / G / FF Weis-Fogh, 1956
2.99   / A / A / FF Weis-Fogh, 1952
2.3-3.5   / A / A / FC Weis-Fogh, 1952
0.5-7   / R / A / WT Zarnack & Wortmann, 1989
3   / A / A / WT Zarnack & Wortmann, 1989
3.7   / A / A / FC Kutsch et al., 1999
Aphididae Myzus persicae 0.3-0.75   / A / A / FF Kennedy & Thomas, 1974
Phorodon humili 7   / A / A / FC Hardie et al., 1996
Mantidae unknown 1.9-3.8   / A / A / FC May, 1999

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Plecoptera

Family Genus species Speed No. / Range / Type / Method Source
Ausroperlidae Acruroperla atra 2.24   / A / G / FF Marden et al., 2000
Tasmoperla thalia 1.56 3 / A / G / FF Marden et al., 2000
Caopniidae various 3.2 (0.29)   / A / G / FF Marden et al., 2000
Capniidae Paracapnia angulata 2.4 1 / A / G / FF Kramer & Marden, 1997
Chloroperlidae Swelta onkos 2.9 1 / A / G / FF Kramer & Marden, 1997
various 6.3 (1.5)   / A / G / FF Marden et al., 2000
Gripopterygidae various 4.4 (1.66)   / A / G / FF Marden et al., 2000
Leuctridae various 4.2 (0.19)   / A / G / FF Marden et al., 2000
Leuctra hippopus 3.7 (0.7) 28 / A / G / FF Kramer & Marden, 1997
Leuctra sibleyi 3.4 (0.54) 4 / A / G / FF Kramer & Marden, 1997
Nemouridae Amphinemura nigritta 2.3 (0.66) 7 / A / G / FF Kramer & Marden, 1997
Ostrocerca spp. 1.6 (0.42) 4 / A / G / FF Kramer & Marden, 1997
Paranemoura perfecta 2.6 (0.54) 8 / A / G / FF Kramer & Marden, 1997
various 3.2 (0.29)   / A / G / FF Marden et al., 2000
Notonemouridae various 6.3 (1.5)   / A / G / FF Marden et al., 2000
Perlodidae Isoperla sp. 2.4 1 / A / G / FF Kramer & Marden, 1997
Perlodidae various 6.3 (1.5)   / A / G / FF Marden et al., 2000
Taeniopterygidae various 3.2 (0.29)   / A / G / FF Marden et al., 2000
Taeniopteryx burski 2.3 (0.36) 5 / A / G / FF Kramer & Marden, 1997
2.9 (0.9) 5 / A / G / FF Marden & Kramer, 1994

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Psocoptera

Family Genus species Speed No. / Range / Type / Method Source
Stenopsocidae Stenopsocus immaculatus 0.85   / A / G / FC Lewis & Taylor, 1967

Editor's Note

This chapter replaces an early version of Chapter 1, by J.H. Byrd. - T.J. Walker, April 2003.

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© Copyright 2003 T.J. Dean. This chapter may be freely reproduced and distributed for noncommercial purposes. For more information on copyright, see Copyright & Permitted Uses.