JOURNAL OF MEDICAL ENTOMOLOGY

J. Med. Entomol. Vol. 17, no. 3: 195-210 31 May 1980

Published bimonthly by Department of Entomology. Bishop Museum, Honolulu, Hawaii, U.S.A. Editorial commiuee: F. J. Radovsky. Executive Editor', JoAnn M. Tenorio, Editor, G. F. Bennett, B. N. Chanimis. M. M. Crystal, R. H. Dadd, R. Damrow, K. C. Emerson, A. Fain, J. L. Grcssitt, H. Hoogstraal, J. Kitzmiller, H, Kurahashi, G, C. laBrecque, F. S. Lukoschus, C. J. Mitchell, M. Nadchatram, W. A. Nelson, M. W. Service, A. Spielman, R. Traub. Devoted to all branches of medical entomology from the world standpoint, including systematics of insects and other arthropods of public health and veterinary significance.

REVIEW ARTICLE!

BIOLOGY AND CONTROL OF THE FACE , (DIPTERA: MUSCIDAE)

By L. G. Pickens and R. W. Miller2

Abstract. The face fly, l\1USca autumnalis, was first discovered fly in North America in 1966 (Smith et al. 1966), in North America in 1952, although it is native to Europe, followed by 5 supplements (Smith & Linsdale Russia, China, North Africa, Korea, Japan, and Iran. During 1967, 1968, 1969, 1971, and Smith & Krancher the past 25 years, the face fly has spread throughout Canada and most of the United States. The face fly is primarily a pest 1974). Since a great deal of the literature concern- of , and its spread in North America was accompanied by ing the face fly has been published since the 1960's, complaints of eye disorders and cattle clustering. Although and since the face fly is still a problem in the USA, there were reports in the literature prior to the 1950's on the biology of the face fly, there were few or no reports on the we attempt, in this paper, to review the research control of this . However, there has since been a consid- literature that has been published on the biology erable amount of research conducted on this pest. This review and control of the face fly from 1916 to 1979, in is an attempt to summarize the published knowledge on the the hope that such a consolidation of research re- biology and control of the face fly. sults will aid current and future researchers who The face fly, Musca autumnalis De Geer, is indig- are concerned with the face fly. enous to Europe, Russia, China, North Africa, Ko- rea, Japan, and Iran (Eltrigham 1916, Hammer BIOLOGY 1941, Kobayashi 1919, Okamoto 1924, Krecob Life cycle 1949, Teskey 1960), It was first discovered in Although the relatively low reproduction rate, North America at Middleton, Nova Scotia in 1952 limited choice of breeding material, and high mor- (Vockeroth 1953), and then spread rapidly west- tality of the immature stages would seem to act to ward across North America in a belt including keep the face fly population relatively low, the rap- southern Canada and the northern United States, id and extensive rate of dispersal of the and In Canada, the face fly spread along rivers and in their habit of feeding on the eyes of cattle make areas where forests and water adjoined agricul- them a pest, even in relatively low numbers. tural land until eventually the insect was found The life cycle of the face fly under pasture con- throughout southern Canada (Depner 1969). In ditions usually requires about 17-21 days (Ham- the United States, the westward and southern mer 1941, Wang 1964) and adult flies live from 20 spread was also rapid; the fly was found in all of to 50 days, depending upon air temperature (Kil- the continental United States except Florida, Lou- lough 1965, Turner & Hair 1966), The males com- isiana, Texas, Alaska, New Mexico, and Arizona mence mating at 2-5 days of age and the females by 1969 (USDA 1969, 1975). at 3-6 days, Males will mate an average of 4 times Much research has been conducted on the biol- and females once, after which sperm are stored in ogy and control of the face fly since its introduction a spermatheca for as long as 28 days (Lodha et al. into North America, and California Vector Views 1970). Most mating occurs near prominent objects published an annotated bibliography of the face in the pasture between 0800 hand 1000 h (Lodha et at 1970) with males taking up positions facing the sun on posts, rocks, etc, and then darting out 'This is the 20th Review Article to appear in the series pub- to grasp females as they fly past (Teskey 1969, lished by the Journal of Medical Entomology. Lodha et at 1970), Males do not go to cattle after 2Livestock Insects Laboratory, Agricultural Environmental Quality Institute, Science and Education Administration, Ag- 2-4 days of age, and 93% of the flies on cattle at ricultural Research, Beltsville, MD 20705, USA. any time are females (Dobson & Matthew 1960, 196 J. Med. Entomol. Vol. 17, no. 3

Teskey 1969). The majority (93%) of the females The flies will often move 0.5 to 11 km over a pe- that are on cattle are present during periods when riod of 3-5 days (Fales et al. 1964, Killough et al. the flies' oocytes are in early stages of maturation 1965, Hansens & Valiela 1967, Ode & Matthysse for each gonotrophic cycle, i.e., at ages 3 and 8 1967). The flies tend to congregate on or near - days (Miller 1967). Hower & Cheng (1972) re- tle at watering places (Pickens et al. 1977), but ported that 5-day-old female flies are 12 times as leave them every evening (Killough et al. 1965) so abundant on cattle as are I-day-old females. Jones that there is a redistribution of the flies every (1963) and Miller & Treece (1968) reported that morning (Engroff et al. 1972). only 4% of the face fly population are on cattle at anyone time. Peak numbers of face flies on cattle Olfactory and visual response occurred between 0700 hand 1300 h and again Both males and females have been reported to between 1500 hand 1700 h (Ode & Matthysse feed on the nectar of umbelliferous flowers (i.e., 1967, Engroff et al. 1972). A peak of male fly ac- wild parsnips and parsley) and on dung, and fe- tivity 3 h after sunrise was reported by Peterson & males commonly feed on blood, sweat, saliva, nasal Meyer (1978) and a peak of mating at 0800 h was mucous, dung, and tears (Teskey 1960, USDA reported by Lodha et al. (1970). Generally, flies 1963). Female flies are attracted to soft cattle feces are more numerous on cattle that are lying down (80-85% moisture content, Bay et al. 1969) lying than on grazing cattle (Benson & Wingo 1963, in full sunlight for up to about 2 h after the feces Hansens & Valiela 1967, Engroff et al. 1972, Par- are dropped. They have also been reported to rish & Gerhardt 1976) and flies are reportedly oviposit in the feces of , swine, or if more common near woods or brush than in open no cattle feces are available (Kobayashi 1919, Vain- pastures (Hammer 1941, Depner 1969). shtein & Rodova 1940, Bay et al. 1968, Barger & Approximately equal numbers of each sex are Anderson 1970). Gravid flies that are seeking sites produced from eggs (Killough & McClellan 1965). in which to oviposit are reportedly most attracted Each female lays an average of 20 eggs in fresh to feces from cattle fed alfalfa diets or grain-sup- bovine feces every 2-8 days during about 21 days, plemented alfalfa diets and are least attracted to for an average of 100 eggs per female lifespan feces from fed grain or corn silage (Treece (Killough & McClellan 1965, Turner & Hair 1966). 1966, Bay et al. 1969). The female flies are also There is a preoviposition period of 5-12 days attracted to amniotic fluids and afterbirth (Teskey (Heller 1976), and ovipositional peaks at 6 and 11 1960). Female response to blood and glucose de- days of adult age (Miller 1967). The eggs usually creases when the females are gravid or in diapause hatch in 18-24 h and the 3 larval instal's last a total (Stoffolano 1968b). Although the flies have been of from 3 to 5 days (Wang 1964), after which lar- reported by some investigators to prefer dark or vae migrate out from the manure pats for a dis- spotted cows to white cows (Hansens & Valiela tance of up to ca 10 m Oones 1969). The pupal 1967, Ode & Matthysse 1967, Engroff et al. 1972), case becomes hardened and white in 24 hand Teskey (1960) observed no fly preference for any adults emerge 7-10 days after pupation (Teskey cattle breed, color, or age. Both male and female 1960). face flies have been attracted to white panels or There has been an observed combined egg and pyramids (Pickens et al. 1977), but only males were larval mortality of 50-90% in the field (Wang attracted to white discs on posts (Peterson & Meyer 1964, Teskey 1969, Valiela 1969, Burton & Turn- 1978). Both sexes are attracted by ultraviolet ra- er 1970) and of 21-26% in the laboratory (Kil- diation of 350-370 nm, but require a higher ra- lough & McClellan 1965). Pupal mortality may also diance than do stable flies, Stomoxys calcitrans (L.), be as high as 56-88% in the field (Valiela 1969). or house flies, Musca domestica L. (Thimijan et al. Nearly all of the mortality of the prepupal stages 1973). in the field is the result of predators attacking eggs or 1st-instal' larvae (Valiela 1969). Sex pheromones Dispersal of the adult flies between pastures or Although face fly females emit a copulatory herds is unaffected by time of day, type of site, fly stimulating pheromone (Chaudhury et al. 1972), age, or fly sex if ambient light and weather con- it does not attract males from more than 4-5 em ditions are optimal (Ode & Matthysse 1967). Flight away (Uebel et al. 1975). Components of the stim- begins when the air temperature is above 17°C ulatory pheromone have been identified as the and the light is suitably intense (Hammer 1941). straight chain monoalkenes (Z)-14-nonacosene, 1980 Pickens & Miller: Biology and control of the face Ay 197

(Z)-13-nonacosene, and (Z)-13-heptacosene. Both 1978, Stoffolano & Matthysse 1967, Valder et at. sexes contain the same saturates and unsaturates, 1969). Oocyte maturation ceases in diapausing fe- but males have a higher proportion of saturates males, hypertrophy of the fat body occurs, and and a lower proportion of active unsaturates than both sexes become photophobic until they become do the females (Uebel et at. 1975). active again in mid- to late April, when mating takes place (Benson & Wingo 1963, Ode & Matthysse Weather effects 1967). Equal numbers of males and females over- In general, face flies are most active on bright, winter at the same site (Ode & Matthysse 1967), warm days when winds are less than 16 km/h. The and nondiapausing adults of both sexes can sur- flies become active when the air temperature is vive constant temperatures of 6 °C for 2-3 weeks above ]6 °C and are most active at temperatures (Morgan & Pickens 1967), while diapausing flies between 25 and 29°C (Hammer 1941, Hansens & can withstand constant temperatures of 5 °C for at Valiela 1967, Ode & Matthysse 1967, Teskey 1969, least 4 months (Caldwell 1975). Peterson & Meyer 1978), providing that the level Caldwell (1975) reported that 70% of the female of incident light exceeds 10,760 Ix in the morning face flies that were collected at Guelph, Ontario or 753.2 Ix at sunset (Killough et at. 1965, Hansens (lat. ca 44°N) were induced to enter diapause when & Valiela 1967, Teskey 1969, Peterson & Meyer they were exposed to daylengths of 14 to 15 h at 1978). Relative humidity apparently has little ef- 16°C, but that only 55% of female flies obtained fect on face fly activity, although rainfall decreases from Nebraska (lat. ca 41ON)would enter diapause activity (Hammer 1941, Matthysse 1961, Engroff under those conditions. The 14- to 15-h photo- et at. 1972). period (morning to evening civil twilight) occurs The flies generally do not enter shelters between 25 August and 12 September at Guelph, or other shaded areas during the summer (Kil- but between 15 August and 8 September in Ne- lough et at. 1965, Ode & Matthysse 1967) although braska, and between 1 August and 1 September in they were reported to be more numerous on cattle Indiana and Kansas. We would therefore expect in the shade of trees than on cattle in the open in that the more southern flies would require a short- some areas (Hansens & Valiela 1967, Parrish & er photoperiod to induce them to enter diapause Gerhardt 1976). They seldom oviposit on fecal than would the northern flies because of the pats that are dropped in heavy shade (Teskey higher temperatures associated with early August. 1969). Winds of less than 16 km/h do not affect fly This seems to be true, as face flies in Indiana and activity, but winds above 16 km/h decrease fly ac- in Kansas enter diapause sites in September (13-h tivity (Hammer 1941, Benson & Wingo 1963, Han- day) (Dobson & Matthew 1960, Valder et al. 1969). sens & Valiela 1967, Engroff et al. 1972). Since the illumination during the civil twilight Face fly populations in North America often intervals (753-2152 Ix) was found to affect the ten- peak in late May, late June, and early August, then dency of the flies to enter diapause (Caldwell & decline in mid- or late August, and have a final Wright 1978), the light intensity required to initi- peak ca 2 weeks later (Caldwell 1975, Ode & Mat- ate diapause must be much lower than the intensity thysse 1967). The first eggs deposited in the spring required to initiate movement in nondiapausing are from females that have overwintered as unin- flies [10,760 Ix reported by Hansens & Valiela seminated adults that have mated prior to leaving (1967)]. the overwintering sites. The flies leave their over- The flies may overwinter in church steeples or wintering sites about the end of April in Missouri, hollow trees and they have been reported to enter New York, and Canada (Benson & Wingo 1963, houses or barns in such numbers that they be- Ode & Matthysse 1967, Teskey 1969), possibly as come a nuisance to homeowners (Eltrigham 1916, a result of being exposed to temperatures of 16- Kearns 1942, Dobson & Matthew 1960, USDA 21°C after a prolonged period (4 months) of very 1961, Benson & Wingo 1963, Strickland et al. low temperatures (Caldwell 1975). 1970). The flies have been reported to overwinter in the same structure year after year, so that some Diapause houses are known as "fly houses" (Caldwell 1975, Overwintering flies enter a true diapause as a Graham-Smith 1919, Matthew et al. 1960). The in- consequence of the exposure of adults that are less fested buildings varied in color and function, but than 2 days old to a combination of low tempera- were usually the tallest buildings in the vicinity tures «16°C) and short days (Caldwell & Wright (Strickland et al. 1970). \98 J. Med. Entomol. Vol. \7, no. 3

Laboratory rearing nificant injury (Shugart et a!. 1979). Herds of cattle Following the development of a successful lab- that have more than 6-10 flies per face have been oratory rearing technique for the face fly (Fales et reported to have a 12% incidence of pinkeye, while al. 1961a), a great deal of work was done to deter- cattle with 16-25 flies per face have a 28% inci- mine the physiological and behavioral character- dence (Cheng 1967) and often stand clustered to- istics of the fly. The adult fly diet originally devel- gether for protection from the flies during the day- oped by Fales was later simplified to consist of time (Teskey 1960, Decker 1961, Hansens & equal parts of table sugar and nonfat dry milk ValieIa 1967). Cattle with ca 50 flies per face have (Turner & Hair 1967). Although at least 1 artificial been reported to have a 60% incidence of pinkeye, larval diet has been attempted (Buletza 1967), compared to 0% infection for cattle with fewer fresh bovine feces is still the only satisfactory larval than 14 per face (Gerhardt & Cook 1976). How- diet. The optimum rearing room conditions were ever, pinkeye may be absent from herds that have reported by Killough & McClellan (1965) and by many face flies or present in herds that have no Turner & Hair (1966) to be 27 ± 1°C, 50 ± 5% face flies (pers. obs.). relative humidity, a 16:8 h light:dark photoperiod, The face fly has also been reported to be a pest and an incident light intensity in the adult colony of bison (Burger & Anderson 1970), (Greg- cage of ca 807-860.8 Ix from cool-white fluores- ory & Wright 1973; J. H. Fales, pers. commun.), cent lights. The adult lifespan in a laboratory (MacNay 1961), and deer (Teskey ranges from 20 days at a constant air temperature 1960), and even of humans (Dobson & Matthew of 32°C to 50 days at 21 °C (Turner & Hair 1966). 1960). Predators and parasites Importance of the face fly Although a variety of organisms have been ob- The spread of the fly in North America was ac- served to attack or affect adult or immature stages companied by farmers' complaints of eye disorders of the face fly (TABLE 1), efforts to use such or- and clustering of their cattle (Dobson & Matthew ganisms as control agents in the field have been 1960, Decker 1961, Benson & Wingo 1963, Treece unsuccessful. Hymenopteran parasitoids, which 1960). There have been reports of decreases in have been used with some degree of success in con- milk production of as much as 25% and decreased trolling other dipteran species, are incapable of weight gain of ca 114 g/day (Peterson & Boreherd- emerging from the calcified face fly puparia, even ing 1962, Hansens 1963), though recent research when a female is able to insert an egg (Burton & data on decreased milk production are lacking. Turner 1968). Intraspecific competition is appar- The face fly has also been found to be a vector ently not a factor in survival (Valiela 1969). of Moraxella bovis, the organism responsible for in- fectious bovine keratoconjunctivitis (pinkeye); lBR CONTROL virus (infectious bovine rhinotracheitis of cattle); When the face fly first became recognized in the Branhamella catarrhalis; Corynebacterium pyogenes; United States as a potential pest of cattle, various and of at least 4 species of eye worms, Thelazia insecticides, repellents, attractants, and combina- rhodesi, T. gulosa, T. skrjabini, and T. lacrymalis (Kre- tions of these were tested for its control. They were cob 1949, Steve & Lilly 1965, Cheng 1967, Stof- applied as wipes, paints, and sprays (Bruce et a!. folano 1971, Weinmann et a!. 1974, Wetzel 1974, 1960, Dobson & Huber 1961, Granett & Hansens Gerhardt et a!. 1976, Geden & Stoffolano 1977). 1961, Matthysse 1961), and later as larvicides fed Although face flies act only as mechanical trans- to cattle to render the feces toxic to developing fly mitters of bacteria [for at least 3 days after contam- larvae (Treece 1962b, 1964, Wallace & Turner ination (Brown & Adkins 1972)], they are true bi- 1964). Although many of the compounds were ef- ological vectors of the Thelazia spp., as the larvae fective in the laboratory or on isolated farms, the are ingested by the fly and then develop in the fly success achieved on commercial farms was erratic to the infective stage, which migrates from the ab- at best. At first the failures were taken as an in- domen to the head of the fly from which they es- dication that many of the insecticides were not cape when the fly is feeding (Krecob 1949). effective against the face fly; however, many sub- Face fly feeding for an equivalent of 33 fly days sequent reports, which will be discussed later, on the eye conjunctiva of cattle can cause severe showed that this was not the case. lesions and has been proposed as an index of sig- A more probable cause was a lack of isolation of 1980 Pickens & Miller: Biology and control of the face fly 199

TABLE I. Organisms that have been reported to attack or affect Musea autumnalis De Geer.

ORGANISM REFERENCES

Acarina Maemeheles muscaedomesticae (Scopoli) Singh et al. 1966 Aves Starlings, Stunltls vulgaris Hammer 1941, Valiela 1969 Boat-Tailed Grackle, Cassidix mexicanus major Fales, J. H. (pers. commun.) Coleoptera Aleoehara bimaculata Gravenhorst Heller 1976, Thomas & Wingo 1968, Kessler & Balsbaugh 1972, Wingo et al. 1967 Aleochara tristis Gravenhorst Heller 1976, Drea 1966, Wingo et al. 1967, Jones 1967 Aphodius jimetarius (L.) Hammer 1941 Lmnellicornia sp. Hammer 1941 Hyponygrus sp. Valie]a ]969 Hister abbreviatus F. Kessler & Balsbaugh 1972 Onthophagus gazella (F.) Lancaster et al. 1976 Philonthus eruenta/us Gmelin Valiela 1969, Kessler & Balsbaugh 1972 P. reetangulus Sharp Kessler & Balsbaugh 1972 Sphaeridium lunatum F. Hammer ]941 S. scarabaeiodes (L.) Rummel 1971 Diptera Sarcophagidae Hammer 1941 Asilidae Pickens (unpubl.) Hymenoptera Aphaertlt palliPes (Say) Kessler & Balsbaugh 1972, Benson & Wingo 1963, BEckie 1961, Hayes & Turner 1971, Houser & Wingo 1967, Garry & Wingo 1971, Rummel 1971, Thomas & Wingo 1968, Wingo 1970, Wylie 1973 Alysia undescribed sp. Thomas & Wingo 1968 Eucoila impatiens (Say) Hair & Turner 1965, Hayes & Turner 1971, Rummel 1971, Thomas & Wingo 1968 Eupteromalus dubius (Ashmead) Wylie 1973 Mellinus sp. Hammer 1941 l\1usidifurax rap/or Girault & Sanders Burton & Turner 1968, Thomas & Wingo 1968, Wylie 1973 Nasonia vitripennis (Walker) Hair & Turner 1965 Spalangia niger Latrielle Burton & Turner 1968, Wylie 1973 Vespa sp. Hammer 1941 Vespula gennanica Schmidtmann 1977 XyaloPhora quinquelineata (Say) Blickle 1961 Nematoda Heterotylenchus autumnalis Nickle Nickle 1967, Stoffolano & Nickle 1966, Kessler & Balsbaugh 1972, Thomas & Puttler 1970, Jones & Perdue [967, Stoffolano 1968a, Treece & Miller 1968, Wright 1972 Ph ycom ycetes Entomophthora sp. Hammer 1941, Gregory & Wright 1973 Entomophthora muscae (Cohn) Fres. Pickens (unpuhl.)

the treated farms (Wallace & Turner 1964). Since in correctly estimating the size of the local face fly face flies disperse rapidly for distances of 3.2 km population. Some entomologists have recommend- or more (Ode & Matthysse 1967, Killough et al. ed counting the flies on an entire animal (Cheng 1965, Pickens et al. 1977), it is difficult to control et al. 1962, Matthysse 1961, 1962, Bruce et al. them by treating only a few farms in an area be- 1960, Decker 1961), while others counted all flies cause of immigrating adults from adjacent untreat- on an animal's head (Dobson & Huber 1961, Gra- ed areas. nett & Hansens 1961, Fales et al. 1961c), and still Another problem in evaluating treatments to others counted the flies on only the face (Holds- control face fly populations has been the difficulty worth 1962). McGuire & Sailer (1962) recom- 200 J. Med. Entomol. Vol. 17, no. 3 mended restricting the count to those flies around days when cattle were sprayed with DDT or me- both eyes of at least 10 animals. Ode & Matthysse thoxychlor. Treece (1960), however, found that (1967) compared the various methods of sampling spraying beef cattle with DDT, toxaphene, or mal- and recommended that the flies be counted on the athion did not give satisfactory control for longer entire head of each of 15 animals. No more than than 1 day; coumaphos and methoxychlor showed 15 animals could be examined because of the mo- more promise. Bruce et at. (1960) also found that bility of both the animals and the flies. Pickens et saturation spraying of cattle with DDT, malathion, al. (1977) reported that sticky traps were a more ronnel, or toxaphene did not achieve a significant sensitive and accurate method of sampling fly pop- level of face fly control. They did, however, find ulations than were counts on animal faces. There that dimethoate and/or dichlorvos, applied in a has generally been little relationship between the syrup bait to the faces of cattle, eliminated most of number of cow fecal pats that contain face fly lar- the face flies for periods up to 5 days. Fales et al. vae and the number of adult flies present on the (1961c), however, reported that baits containing cattle (Wallace & Turner 1964, Hansens 1963). malathion, methoxychlor, trichlorfon, or dichlor- vos failed to give satisfactory control of the face Susceptibility of face flies to insecticides fly, although the latter 2 compounds were better Treece (1961) showed that the face fly was more than the other 2. Sprays containing barthrin [6- susceptible than the house fly to DDT and as sus- chloro-l,3-benzodioxol-5-yl 2,2-dimethyl 3-(2- ceptible to parathion. Bodenstein & Fales (1962) methyl-l-propenyl)cyclopropanecarboxylate] and compared the susceptibility of face flies and house piperonyl butoxide that were applied to the head flies to 21 different insecticides, which included or head and front quarters of dairy and beef cattle organochlorines, organophosphates, and carba- were moderately effective into the 2nd day after mates. They concluded that the face fly was more treatment. susceptible than the house fly to all the insecticides, It was reported in the early 1960's that naled with the possible exception of endrin, toxaphene applied as a water-dispersed spray (Granett & and terpene polychlorinates [65% chlorine] (5tro- Hansens 1961) or as a bait spray (Fales et al. 1961b) bane®). Turner & Wang (1964) tested the same 3 gave a moderate degree of control of the face fly classes of insecticides and concluded that the face and was more effective than other insecticides fly was susceptible to a wide range of organic in- tested. At about the same time, other workers secticides. Roussell (1965) also found a high degree (Matthysse 1961, Treece 1962a) reported that di- of susceptibility to the same 3 classes of insecticides chlorvos applied as a sugar bait directly to the faces in both laboratory-reared and field-collected of cattle offered protection from the face fly for a strains of face flies. day or more. Holdsworth (1962) concluded that Bodenstein et at. (1971) evaluated 281 insecti- the conscientious daily application of dichlorvos cides from 1961 to 1969 and reported that 24 gave sugar bait could greatly alleviate face fly annoy- 100% control of the face fly with a residue of 0.312 ance, but that irregular use was not satisfactory. mg/929 cm2 or less. Fales & Bodenstein (1966) and Dorsey (1966) tested dichlorvos baits with and Fales et al. (1971) reported that the synthetic py- without the addition of 2 other insecticides, azin- rethroid insecticides tetramethrin and d-trans-al- phosmethyl or phosmet for fly control on horses. lethrin were more effective than DDT against the The combination product gave the best results, but face fly. only fairly good protection for 2 h and limited pro- Robinson et al. (1975) tested several organo- tection for up to 24 h. This was considered neither phosphorus insecticides, an organochlorine (me- practical nor effective. Dorsey (1966) did, how- thoxychlor), and a synthetic pyrethroid (permeth- ever, show that dichlorvos-strand halters on horses rin) against laboratory-reared face flies originally provided effective relief from face fly annoyance collected in Mississippi. Naled and permethrin within the area protected by the strands. were the most effective, with methoxychlor the Granett & Hansens (1961) found that a stabi- least. Lancaster & Simco (1975) also reported that lized dust formulation of dichlorvos was more ef- the face fly was quite susceptible to permethrin. fective for face fly control than a water-base spray or as paint or liquid treatment applied directly to Hand application of insecticides to cattle the faces of cattle. Even so, the dust only markedly Dobson & Huber (1961) reported that control reduced face fly annoyance for 6-9 h after appli- of the face fly was excellent and lasted for 10-14 cation. They also reported that naled, applied as 1980 Pickens & Miller: Biology and control of the face fly 201 a stabilized dust or spray, provided 50-60% re- horn fly, Haernatobia irritans (L.), control (Rogoff duction in face fly populations on the day of ap- & Moxon 1952) and had been in use for this plication, but caused some discomfort to the ani- species, it was logical ta investigate them for face mals and the applicator. Other insecticides were fly control. also tested with varying, but usually limited, de- Dobson & Huber (1961) used cable-rubber de- grees of success. vices and reported that populations of the face fly Dorsey et al. (1962) found organophosphorus were adequately reduced by methoxychlor, toxa- sprays to be generally ineffective against the face phene, DDT, and ronnel. Dimethoate and dichlor- fly; however, diazinon and dimethoate offered fair vos were not effective in reducing face fly popu- to good protection for 7-14 days. Dust applications lations. of some insecticides also offered some degree of Tests were conducted in Georgia (Roberts 1963, protection for about the same length of time. They Roberts et al. 1963) to compare various rubbing reported that several organophosphorus insecti- devices treated with toxaphene, carbaryl, and or- cides in smear-type formulations afforded cattle ganophosphorus insecticides. Although all insec- limited protection against the face fly for about 7 ticides in rubbing devices tested gave good ta ex- days. cellent control of the horn fly, the maximum Knapp (1962) reported inconsistent control of degree of face fly control achieved was 38%. In a the face fly with a formulation containing ronnel, later series of trials with dust bags, Roberts (1965) pyrethrins, and piperonyl butoxide; however, pop- reported 38% and 85% control of the face fly with ulations were very low in that study. methoxychlor and carbaryl, respectively. Hansens & Granett (1963) found crotoxyphos to Turner (1965) tested self-applicating dust dis- be superior to any compound that they had pre- pensers for the cantrol of face flies on a 81O-ha viously tested against the face fly (Granett & Han- farm that had cattle in 23 separate pastures. Var- sens 1961, Granett et al. 1962). Also effective were ious organophosphorus insecticides and a propri- chlorfenvinphos, dimetilan, and Pyramat (6 meth- etary product containing pyrethrins were used in yl-2 -propy 1-4-pyri mid in yI d imeth ylcarba mate). the dispensers. Significant, but not complete, re- These compounds were all reported to be more ductions of face flies were obtained. effective than several pyrethrins, carbamates, and Hair & Adkins (1965) reported that bags con- organophosphates, including dichlorvos, previ- taining dusts or wettable powder formulations of ously reported to be one of the more promising insecticides were more effective than were back- compounds for face fly control. rubbers for controlling face Aies. They reported DeFoliart (1963) also found that crotoxyphos that dust bags containing dimetilan, azinphos- was effective against face flies when sprayed week- methyl, or toxaphene + DDT gave excellent and ly and starting before flies were numerous. Oil- consistent control. Dorsey et al. (1966) reported and water-based formulations of this insecticide effective (up to 60% reduction) of face flies with provided good control of face flies for 28 hand dust bags and backrubbers containing various in- significant reductions of fly populations for 52-76 secticides. Azinphosmethyl and propoxur were the h following spray applications (Cheng et al. 1965). most effective. Fales et al. (1968) also reported positive results in Adkins & Seawright (1967) described a relatively terms of face fly control with crotoxyphos against inexpensive dusting station that they said gave con- the face fly on both beef and dairy cattle. The sistent control of the face fly if cattle were forced degree of efficacy was increased when sugar was to pass through it not less than once every 2 days. added to the spray. These same workers (Seawright & Adkins 1968) Wallace & Turner (1962) and Ode & Matthysse tested the stations in the field and reported reduc- (1964a) evaluated many of the insecticides previ- tions in face fly populations from 54% to 85% ously mentioned as dusts, sprays, or baits in field when the dust bags contained dimetilan, ronnel, experiments under farm conditions and found coumaphos, malathion, or stirofos. The same none of them to be particularly effective for con- group (Poindexter & Adkins 1970) reported good trolling face flies. to excellent control of the face fly in a field trial with dust bags containing coumaphos, methoxy- Self-treating devices for cattle chlor, crotoxyphos, toxaphene + DDT + lindane, Since self-treatment rubbing devices had been or DDT + methoxychlor. Fales et al. (1968) found to be an effective and economical means of showed an initial reduction in face flies when cattle 202 J. Med. Enlomol. Vol. 17, no. 3 were placed in a pasture with a dust bag containing have been screened in the laboratory for repellen- 3% crotoxyphos. cy to face flies. Frishman & Matthysse (1966) re- Wrich (1970) reported that in South Dakota cou- ported that face flies, as well as house flies, were maphos and fenthion in dust bags and in back rub- strongly repelled by short, straight-chain aliphatic bers aided in reducing face fly populations. Com- esters. Commercial fly repellents that showed re- plete control of the face fly was obtained in 1 pellent properties in the laboratory, but were not experiment by using a tank-type backrubber con- as effective as the simple esters, included MGK taining 1% fenthion. In contrast, Kessler & Berndt R-1207, MGK R-l1 [1,5a,6,9,9a,9b-hexahydro- (1971), also from South Dakota, reported that nei- 4a(4H)-dibenzofurancarboxaldehyde], MGK-874 ther dust bags containing crotoxyphos nor back- [2-(octylthio)ethanol], MGK 326 (hexanediol), di- rubbers containing toxaphene and motor oil sig- butyl phthalate, and butoxy polypropylene glycol. nificantly reduced face fly numbers below those in When these compounds were tested in the field, a check herd. Ronald & Wingo (1973) also report- the most effective were MGK R-1207, MGK R-ll, ed no control of the face fly when cattle were ex- MGK 326, dibutyl phthalate, hexanediol, and bu- posed to toxaphene-treated backrubbers. toxy polypropylene glycol. The highly volatile re- Gel bags containing an encapsulated formula- pellents were not effective in the field. In these tion of dichlorvos and stirofos were no more ef- field experiments, it was observed that when di- fective for face fly control than a standard cou- chlorvos was added to corn syrup, flies were re- maphos dust bag (Boxler et al. 1977). A 2nd pelled. This confirmed an earlier observation by experiment with a dichlorvos gel bag also did not Decker (1961). provide control of the face fly. Bodenstein et al. (1970) screened 75 compounds Another self-applicatory approach was investi- for repellency to the face fly. Fifteen of the com- gated by Cheng et al. (1962), who reported that a pounds were as effective as MGK 326, the stan- 1% dimethoate spray applied with an electric eye- dard used for comparison, when they were tested controlled sprayer or a photostatic aerosol appli- at a concentration of 1.24% or less. The repellent cator gave moderate control of the face fly. activity mainly occurred with 2 general types of compounds, sulfides and sulfoxides, and N,N -di- Face fly repellents substituted amides. Many of the workers cited in the previous sec- tions evaluated repellent formulations alone or in Face fly attractants combination with insecticides (Bruce et al. 1960, In the field tests in which insecticides were hand- Treece 1960, Granett & Hansens 1961, Fales et al. applied to the cows, many were mixed with molas- 1961c, Granett et al. 1962, Cheng et al. 1962, ses baits that acted as attractants (Bruce et al. 1960, DeFoliart 1963, Hansens & Granett 1963, Ode & Fales et al. 1961c, Decker 1961, Matthysse 1961, Matthysse 1964a). Compounds tested included Ta- Treece 1962a, Holdsworth 1962, Benson & Wingo batrex (dibutyl succinate) MGK R-264 [N-(2-ethyl- 1963). Dorsey (1966) also reported on adding in- hexyl)bicyclo(2.2.1 )hept -5-ene-2 ,3-dicarboximide], secticide-sugar baits (syrup, malt, molasses, or hon- MGK R-326 (dipropyl pyridine-2.5-dicarboxylate), ey) to the faces of horses. Benson & Wingo (1963) MGK R-1207 (3-chloropropyloctyl sulfoxide), and reported that honey was 60% more effective than butoxy polypropylene glycol. The results found in syrup for face flies, while Decker (1961) reported these tests were reviewed in detail by Frishman & that the simplest and most effective bait he tested Matthysse (1966). In general, the conclusions consisted of 75% corn syrup and 25% water. drawn were that the repellents tested under field Frishman & Matthysse (1966) field-tested var- conditions gave from mediocre to adequate con- ious compounds they had shown to be attractive to trol. None of the repellents, however, were re- face flies in the laboratory by impregnating blotter ported to last for over 1 day in the field. paper and placing it above the entrance to a "cow Corba (1970), however, reported that an oint- tunnel." Under these conditions, honey, corn syr- ment made of 30% deet repelled face flies on pas- up, sucrose, and citrated bovine blood attracted tured cattle for 3 days. The ortho isomer of deet, the greatest number of face flies. Hammer (1941) dimethyl phthalate, and pyrethrin were also tested, and Treece (1961) had previously reported that but were less effective. face flies were highly attracted to blood. In addition to field experiments to test commer- Dorsey (1968) tested face fly attractants in the cial formulations of repellents, various compounds field by applying them directly to steers or putting 1980 Pickens & Miller: Biology and control of the face fly 203 them in petri dishes placed on elevated platforms those compounds against the face fly. They dem- along the edge of the pasture. The attractants were onstrated marked sterility effects when face fly pu- more effective when applied on the animals than pae were dipped in a solution of apholate or when in the petri dishes. When applied to animals, the adults received apholate or tepa in their diet. most effective materials in 3 different years con- In further work with apholate, Hair & Turner sisted of gut slime, placenta powder, and equal vol- (1966) showed that the sterility effects were greater umes of blood hydrolysate and dimethyl disulfide. when sexually mature females were fed the com- It has been reported that dry ice increases the pound (0.5% of diet) than when it was fed to males. catch of face flies in plastic cone-type traps (Cald- The difference between sexes was not evident well 1975) and in a malaise trap (Pechuman & Bur- when newly emerged flies were fed apholate. ton 1969). Zapanta & Wingo (1968) tested heliotrine, a pyr- Color attraction and preferences of the face fly rolizidine alkaloid, as a sterility agent for face flies. were discussed earlier in this review. Adult face flies treated at the egg or larval stages Since even the best repellents and attractants with 32 g of heliotrine/g of dry manure showed a tested to date have only a short duration of effec- reduction of 52.4% in oviposition and 62.5% in tiveness when applied to cattle in the field, it is egg viability. Puparium production was also de- doubtful that either of these classes of compounds creased, as was formation of mature eggs in ova- will be used on a commercial basis to protect cattle nes. from the face fly. Kaur & Steve (1969) reported that male and fe- male face flies were effectively sterilized when fed Ultra-law-volume (ULV) application of insecticides ad libitum for 6 days on a 0.1 % concentration of to cattle hempa or metepa. Sterility was maintained for 28 Dobson & Sanders (1965) reported on an ex- days in both sexes. In a later study, Kaur & Went- periment in which beef cattle were sprayed with. worth (1972) showed that the same compound technical malathion at a rate of 226.8 g/O.4 ha from sterilized 2- to 3-day-old pupae that were dipped a spray plane. Control of both face flies and horn into a 4.5% acetone solution of either compound. flies was very effective for a few days and adequate Face fly larvae, however, were not sterilized at a for 1 week. Knapp (1966), using similar techniques 0.5% concentration of the compounds in bovine to apply trichlorfon at a rate of 85.0 g/O.4 ha, re- manure. Concentrations higher than this killed the ported excellent control of the face fly in 2 herds larvae. for up to 24 h. Counts were not made after this Oil of Sterculia foetida L. added to dry face fly time. In a follow-up trial with trichlorfon (Knapp diet at a concentration of 9.2% (wtlwt) caused ste- 1967), very little control of the face fly was ob- rility in female flies that was reversible when the tained, which he attributed to immigration from flies were returned to an untreated diet (Lang & a nearby untreated herd. Treece 1971). These same workers (Lang & Kantack et al. (1967) claimed that immigration Treece 1972) reported similar effects when adult of the face fly from adjacent herds did not occur flies were fed a diet containing 1% boric acid. in their experiment when UL V applications of Pickens & DeMilo (1977) reported that when technical malathion (170.1 g/0.4 ha) were made 4 either sex of face flies was fed diflubenzuron in the times during a I-month period. Face fly popula- adult diet or exposed to it by surface contact, egg tions on the treated herds were reduced from 20 hatch was severely inhibited. Three closely related to 5 flies/face. In a later experiment, the South analogues of diflubenzuron also inhibited hatch, Dakota workers Del Fosse & Balsbaugh (1974) re- but none was significantly more effective than di- ported reductions in face fly populations with mal- flubenzuron. Although the mode of action of this athion, stirofos or a mixture of dichlorvos and sti- class of compounds is different than that of the rofos (1:4, Ravap®). traditional chemosterilants, the end result is the same, in that a new generation of flies is not pro- Sterilization techniques for face.fly control duced. The use of chemosterilants has also been inves- Radiosterilization of the face fly has also been tigated in attempts to control the face fly. La- studied. Gregory & Wright (1973) reported that a Brecque (1961) reported that tepa and apholate dose of 2.5 krad of gamma-ray radiation to 5-day- sterilized both sexes of the house fly when fed in old face fly pupae produced complete sterility in the adult diet, and Hair & Adkins (1964) evaluated both sexes. 204 J. Med. Entomol. Vol. 17, no. 3

The face fly, therefore, can be sterilized by was no difference in adult populations of face flies either chemosterilants or gamma radiation, but between the treated and control herds. They con- there has been no research to test this control tech- cluded that the reason was probably migration of nique under field conditions. flies from untreated to treated fields. This conclu- sion was verified in a later experiment (Wallace & Feed additives for face fly control Turner 1964) when 1600 cattle in 23 pastures were fed a mineral salt mixture containing 5.5% ronnel. Feed additives, like other control methods tested Although larval control ranged from 88% to 99%, against the face fly when it first became a problem adult control ranged from 0 to 47%. Again, it was in the United States, were considered because they concluded that immigration of flies from untreat- had previously been reported to offer promise of ed herds was the reason for the poor adult control. controlling other fly pests of cattle (Eddy & Roth Knapp (1965a) also obtained poor control of 1961). adult face flies on cattle offered 5.5% ronnel min- The first report was that of Anthony et al. eral blocks or granules. He concluded that the rea- (1961). They found that the addition of couma- son for the poor control was that not all the cattle phos, Bayer 22408 [2-([diethoxyphosphino- consumed sufficient ronnel to give complete larval thioyl]oxy)-IH -benz(d, e)isoquinoline-l,3 (2H)- control, but did not dismiss the possibility that im- dione], or ronnel completely inhibited development migration of adult face flies into the test area could of face fly larvae when they were fed at levels of also have caused some of the problem. On the oth- 1.0, 0.5, and 2.5 mg compound/kg body wt per er hand, Ronald & Wingo (1973) reported that day, respectively. Subsequently, many organo- a 5.5% ronnel supplement program significantly phosphorus insecticides were screened as feed ad- reduced the seasonal mean number of face flies ditives for control of face fly larvae (Treece 1962b, in each of 2 years on cattle in central Missouri. 1964, Ode & Matthysse 1964b). In general, as re- Jones & Medley (1963) fed cattle coumaphos at viewed by Miller(1970), the more promising com- a rate of 0.5 mg/kg body wt/day or sprayed a 6.48- pounds were those originally mentioned by An- ha pasture with 37 g of coumaphos weekly for 6 thony et al. (1961) with the addition of stirofos weeks. The feces from these cattle were sufficiently (Rabon®). Because of these promising results, ron- toxic to kill face fly larvae, but counts of adults on nel, coumaphos, and stirofos have been tested ex- the cattle were not reduced. They also concluded tensively under field conditions. that migration from untreated herds was the rea- Another compound, dichlorvos, was reported by son for the lack of adult control. Knapp (1965b) Treece (1962b) not to be effective against face fly obtained similar results when 2% coumaphos salt larvae when fed to cattle at a level of 5 mg/kg body was offered to pastured steer calves, as did Dorsey wt/day. Pitts & Hopkins (1964), however, showed et al. (1966) when they fed cattle a coumaphos-salt that a polyvinyl chloride (PVC) resin formulation mixture at a rate of 2 mg active ingredient (AI)/kg of dichlorvos fed at a level as low as 0.5 mg/kg body wt per day. body wt/day gave excellent control of house fly lar- Miller & Pickens (1975) showed ca 50% reduc- vae and also reported that face fly larvae were tion in face fly population from 1 June to 30 Sept. twice as susceptible as house fly larvae to this com- on dairy heifers and dry cows that were fed a con- pound. Lloyd & Matthysse (1970) showed that di- centrate containing 154 ppm (increased by 50% 6 chlorvos, or diazinon, incorporated into PVC and weeks after the start of the experiment) of stirofos fed to cattle at a rate of 0.5 mg/kg body wt per day (calculated to provide 1.5 or 2.25 mg stirofos/kg effectively controlled face fly larvae in manure body wt per day). Gerhardt & Cook (1976) fed a from the insecticide-fed cows. When PVC-dichlor- 1.94% stirofos salt supplement to ca 1200 pastured vos was fed to a dairy herd at a level of 0.25 mg/ beef cattle in an isolated area of Tennessee. From kg body wt per day for 2 months, little control of 1 May to 31 August, there were 52% fewer face face fly larvae occurred. Raising the rate to 0.5 mg flies on the treated cattle than on the controls. Dur- did cause a high degree of larval mortality. ing the time of the highest face fly populations Wallace & Turner (1962) placed a ronnel-salt mixture in the salt boxes of 72 pastured cattle. The Uune and July), the reduction was 72%. This com- daily rate of consumption of the ronnel was cal- pared to a 66% reduction in another herd where culated to be 3 mg/kg body wt per day. Although cattle were sprayed and allowed access to dust bags larval counts indicated a 58-90% reduction, there and backrubbers. 1980 Pickens & Miller: Biology and control of the face fly 205

Over the past several years, the recently devel- In addition to the organophosphorus insecti- oped insect growth regulators have been tested as cides and insect growth regulators that have been feed additives for face fly control. Miller & Uebel tested against the face fly, certain bacterial and bo- (1974) reported that methoprene fed to cattle at tanical insecticides and xanthine dyes have also a level of 1 mg/kg body wt per day gave essentially been evaluated. complete control of the face fly in small-scale tests. Cantwell et al. (1964) placed last-instar face fly Ro 20-3600 [(£)-4-([6,7 -epoxy-3, 7-dimethyl-2-no- larvae into cattle feces that contained a superna- nenyl]oxy)-1,2-(methylenedioxy)benzene], while tant liquid from a commercial product of Bacillus not as effective as methoprene, did give some face thuringiensis Berliner at levels of I, 2, 3, or 4 ppt. fly control at levels ranging between 1 and 5 mg/ The LD50 was determined to be 3.6 ppt, 10 times kg body wt per day; and 10 mg of Ro 20-3600/kg that for the house fly. Yendol & Miller (1967), in body wt per day gave complete control. tests similar to those of Cantwell et at. (1964), de-

Miller et al. (1978) tested methoprene as a feed termined t.hat the LD50 values for the supernatants additive for face fly control under pasture condi- of 2 commercial preparations of B. thuringiensis tions. In 1975, when methoprene was fed in a were 2.06 and 9.46 ppt. They also reported that block formulation at an average rate of 0.5 mg AI/ when B. thuringiensis was fed to a cow at a rate of kg body wt per day, bioassays and counts of adult 36 mg/kg body wt per day, manure from the cow flies on cattle indicated that face fly populations caused up to 100% mortality of 12-h-old face fly were reduced. In 1976, when methoprene was fed larvae seeded into the feces. to similar cattle in a loose mineral supplement at Hower & Cheng (1968) fed dairy cattle B. thu- an average rate of 0.07 or 0.13 mg AI/kg body wt ringiensis at levels of 185 or 370 mg/kg body wt per per day, 45-90% of face fly pupae did not produce day. Inhibition of development of face fly larvae adults, but counts of adults on cattle were not seeded into the manure from t.hese cows averaged reduced markedly. 84.9% and 99.6%, respectively. Ode & Matthysse Schwarz et al. (1974) synthesized and tested 6 (1964b) reported that while B. thuringiensis fed at arylterpenoid compounds that are structurally levels up to 30 mg/kg body wt per day was inef- somewhat similar to methoprene. The most active fective against the face fly, barthrin, dimet.hrin, compound fed to cattle was AI3-36093 [2-ethoxy- and the 3,4 isomer of dimethrin fed at a level of 9-(p-isopropyl phen yl)-2,6-dimethylnonane], which 10 mg/kg body wt per day were very effective. gave 100% inhibition of face fly development when Mississippi workers (Anonymous 1977) fed fed at a level of 0.25 mg/kg body wt per day. Fur- erythrosin B and rose bengal dyes to cattle and ther work with these compounds showed that the found, depending upon the level fed, that the dye methoxy homologue of this compound, AI3- could kill larvae, pupae or adults emerging from 36206, was as active as A13-36093 as a cattle feed the pupae. additive against the face fly (Miller & Schwarz The feed-additive approach for control of the 1975, unpubl. data). face fly appears to be sound since the fly breeds Miller (1974) reported that the insect growth almost exclusively in fresh cattle manure. How- regulator diflubenzuron fed to cattle at a rate of ever, even though several compounds have been 0.25 mg/kg body wt per day killed 90% of the face used successfully to eliminate the development of fly larvae seeded into the feces and no adults immature stages of the fly in the manure, control ec\osed from the 10% that did pupate. of adult flies on the faces of cattle has been from In experiments in our laboratory (Miller, 1978, good to none depending primarily on the degree unpubt. data) we have evaluated both methoprene of isolation from surrounding untreated areas. and diflubenzuron incorporated into sustained-re- Since face flies disperse readily for distances of 3 lease bolus formulat.ions similar to those described km or more, it appears that for a successful feed by Miller et at. (1977). These boluses, which con- additive face fly control program to have any tained 10% active ingredient, gave essentially com- chance of success it should only be attempted on plete control of the face fly larvae in laboratory an area-wide basis. bioassays over 20 weeks. On the basis of the Another, probably somewhat less significant, amounts the boluses eroded and the weights of the factor for the lack of face fly control using this cattle, the bolus method of administering the insect method under field situations can be uneven con- growth regulators appears to be more efficient sumption of the additive when administered via than feeding the compounds. self-fed mineral supplements. 206 J. Med. Entomol. Vol. 17, no. 3

CONCLUSIONS Anonymous. 1977. Face flies-researchers study pesticide po- tential of dyes. Miss. Agric. For. Exp. SIn. Res. Highlights In the relatively short period of time the face fly 40(8): 7-8. has been in North America, it has become recog- Anthony, D. W., N. W. Hooven &: O. Bodenstein. 1961. Tox- nized as one of the major fly pests of cattle and icity to face fly and house fly larvae of feces from insecti- also of horses. Although the face fly is quite sus- cide-fed cattle. J. Econ. Entomol. 54: 406-08. Bay, D. E., C. W. Pitts &: G. M. Ward. 1968. Oviposition and ceptible to insecticides commonly used to control development of the face flies in feces of six species of an- other fly pests of cattle, small-scale field trials, imals. J. Econ. Entomol. 61: 1733-36. which have been conducted in attempts to control 1969. Influence of moisture content of bovine feces on ovi- the face fly, have been generally unsuccessful. position and development of the face fly. I Econ. Entolnol. 62: 41-44. Such failures are due primarily to a lack of isola- Benson, O. L. &: C. W. Wingo. 1963. Investigations of the face tion of treated farms from untreated farms, from fly in Missouri. J. Econ. Entomol. 56: 251-58. which face flies disperse. The dispersal habit, and Btickle, R. L. 1961. Parasites of the face fly in New Hamp- the fly's reluctance to enter buildings except to shire. I Econ. Entolnol. 54: 802. Bodenstein, O. F. &: J. H. Fales. 1962. Residual tests on face overwinter, combined with the presence of only a flies. Soap Chell!. Spec. 38: 125-28. small proportion of the flies on cattle at anyone Bodenstein, O. F., J. H. Fales, R. P. Hall, G. D. Mills, Jr &: R. time, make the face fly one of the most difficult of L. Walker. 1971. Laboratory evaluations of candidate in- the fly pests of cattle to control. secticide residues against face flies and DDT-resistant house flies, 1961-69. ARS-USDA Prod. Res. Rep., Washing- Therefore, future research in controlling the ton, D.C. No. 132.41 p. face fly should attempt to exploit weak points in Bodenstein, O. F., J. H. Fales &: R. L. Walker. 1970. Labo- the biology of this species by using an integrated ratory evaluations of materials as repellents for the face approach. Since the face fly breeds almost exclu- fly, 1962-69. J. Econ. Entomol. 63: 1752-55. Boxler, D. J., J. I. Shugart &: J. B. Campbell. 1977. Use of gel sively in fresh cow feces, the use of feed additives encapsulated slow-release insecticides for control of the face (insecticides or insect growth regulators) should be fly and horn fly on cattle in Nebraska. I Kam. Entomol. Soc. an effective control method if used on an area- 50: 576. wide basis. New and novel ways to administer the Brown, J. F. &: T. R. Adkins. 1972. Relationship of feeding activity of face fly (Musca autumnalis De Geer) to produc- compounds to cattle, such as in a bolus formula- tion of keratoconjunctivitis. Am. J. Vet. Res. 33: 2551-55. tion, should be further investigated. It is impor- Bruce, W. N., S. Moore, III &: G. C. Decker. 1960. Face fly tant, however, that compounds administered to control. I Econ. Entoll!ol. 53: 450-51. Buletza, G. F., Jr. 1967. Nutritional studies on oligidic diets cattle for the control of fly larvae not adversely for the larval stage of the face fly, Musca autumnalis. M.S. affect nontarget species. thesis, Clemson Univ., Clemson, S.c. 45 p. In addition to larval control, methods to elimi- Burger, J. F. &: J. R. Anderson. 1970. Association of the face nate adult face flies should be further investigated fly, Musca autumnalis, with bison in North America. Ann. Entoll!ol. Soc. Am. 63: 655-59. on an area-wide basis. The methods might include Burton, R. P. &: E. C. Turner, Jr. 1968. Laboratory propa- forced-use dust bags or face rubbers, insecticide- gation of Muscidifurax raptor on face fly pupae. J. Econ. impregnated ear tags, and the use of white panels Entomol. 61: 1380-83. to attract adult flies. Investigations should be made 1970. Mortality in field populations of face fly larvae and pupae. J. Econ. Entornol. 63: 1592-94. of ways to utilize these panels to kill the attracted Caldwell, E. T. N. 1975. Diapause and overwintering of the flies with insecticides, to sterilize them with che- face fly, Musca autumnalis De Geer (Diptera: Muscidae), mosterilants, or to capture them mechanically. near Guelph, Ontario. M.S. thesis, Univ. of Guelph. 127 p. Caldwell, E. T. N. &: R. E. Wright. 1978. Induction and ter- Another weak spot in the biology of face flies, mination of diapause in the face fly, Musca auturnnalis (Dip- which deserves investigation, is their habit of over- tera: Muscidae), in the laboratory. Can. Ent01nol. 110: 617- wintering as unmated adults. Research is needed 22. on this stage of the flies' life to find out what makes Cantwell, G. E., A. M. HeimpeI &: M. J. Thompson. 1964. The production of an exotoxin by various crystal-forming them go to certain sites, and to determine the nature bacteria related to Bacillus thuringiensis var. thuringiensis of their migration patterns when they come out in Berliner. J. Insect Pathol. 6: 466-80. the spring. If these facets of the flies' behavior Chaudhury, M. F. B., H. J. Ball &: C. M. Jones. 1972. A sex were better understood, methods could.be devised pheromone of the female face fly, Musca autumnalis, and its role in sexual behavior. Ann. Entomol. Soc. Am. 65: 607- to reduce fly numbers at times when the popula- 12. tions are low. Cheng, T. H. 1967. Frequence of pinkeye incidence in cattle in relation to face fly abundance. j. Econ. Entomol. 60: 598- LITERATURE CITED 99. Adkins, T. R.,Jr &: J. A. Seawright. 1967. A simplified dusting Cheng, T. H., D. E. H. Frear &: H. F. Enos, Jr. 1962. The use station to control face flies and horn flies on cattle. J. Econ. of spray and aerosol formulations containing R-1207 and Entomol. 60: 864-68. dimethoate for fly control on dairy cattle and the deter- 1980 Pickens & Miller: Biology and control of the face fly 207

mination of dimethoate residues in milk. I Econ. Entornol. of the face fly, Musca auturnnalis, and the house fly, Musca 55: 39-43. domestica L. Cornell Univ. Exp. Sin. Mem. 394. 91 p. Cheng, T. H., A. A. Hower & R. K. Sprenkel. 1965. Oil-based Garry, C. E. & C. W. Wingo. 1971. Factors affecting parasitism and water-based ciodrin sprays for fly control on dairy cat- of the face fly by APhaereta pallipes in laboratory studies. I t1e.I Econ. Entomol. 58: 910-13. Econ. Entornol. 64: 104-07. Corba, J. ]970. The use of repellents for the protection of the Geden, C. J. & J. G. Stoffolano. 1977. Musca autumnalis (De eye of cattle against intermediary hosts of Thelazia spp. Vet. Geer) (Diptera: Muscidae) as a vector of Thelazia sp. (Bose) Cas. 13: 59-63. (Nematoda: Filaroidea) in Massachusetts. I N.Y. Entomol. Decker, G. C. ]96]. Face fly problems and control. Soap Chern. Soc. 85: 175. Spec. 37: 101, 103, 105-07. Gerhardt, R. R. & C. W. Cook. 1976. A large-scale test of an DeFoliart, G. R. 1963. Preventive spraying schedules for dairy oral insecticide for controlling face flies. Tenn. Farm Home farm fly control. I Econ. Entornol. 56: 649-54. Sci. Prog. Rep. 99: 9-10. Del Fosse, E. S. & E. U. Balsbaugh,Jr. 1974. Effects ofULV Gerhardt, R. R., G. Parrish, R. Q. Snyder & R. D. Freelana. organophosphates on horn flies and face flies of cattle, and 1976. Incidence of pinkeye, Moraxella bovis, in relation to on the bovine coprocoenosis. Environ. Entornol. 3: 919-22. face fly control. Tenn. Farm Home Sci. Prog. Rep. 97: 14- Depner, K. R. 1969. Distribution of the face fly in western 15. Canada and the relation between its environment and pop- Graham-Smith, G. S. 1919. Further observations on the habits ulation density. Can. £ntornol. 101: 97-100. and parasites of common flies. Parasitology ll: 347-84. Dobson, R. C. & D. A. Huber. 1961. Control of face flies, Granett, P. & E. J. Hansens. 1961. Tests against face flies on Musca au/urnnalis, on beef cattle in Indiana. I Econ. Ento- cattle in New Jersey during 1960.I Econ. Entornul. 54: 562- mol. 54: 434-36. 66. Dobson, R. C. & D. L. Matthew. 1960. Field observations of Granett, P., E.J. Hansens & A.J. Forgash. 1962. Tests against face fly (Musca aulwnnalis De Geer) in Indiana. Proc. In- face flies on cattle in New Jersey during 1961. I Econ. diana Acad. Sci. 70: 152-53. Entorno/. 55: 655-59. Dobson, R. C. & D. P. Sanders. 1965. Low-volume, high-con- Gregory, D. & R. E. Wright. 1973. Irradiation of the female centration spraying for horn fly and face fly control on face fly, Musca auturnnalis De Geer, parasitized by the nem- beef cattle. I Econ. Entornol. 58: 379. atode, Heterotylenchus autumnalis Nickle. Proc. Entornol. Soc. Dorsey, C. K. 1966. Face fly control experiments on quarter Onto 103: 104-07. horses-I 962-64. I Econ. Entornol. 59: 86-88. Hair, J. A. & T. R. Adkins, Jr. 1964. Sterilization of the face 1968. Field experiments with attractants for the face fly. I fly, Musca auturnnalis, with apholate and tepa.I Econ. Ento- Econ. Entomol. 61: 1695-96. rno/. 57: 586-89. Dorsey, C. K., J. O. Heishmann & C. J. Cunningham. 1966. 1965. Dusting stations and cable backrubbers as self-appli- Face fly and horn fly control on cattle. I Econ. Entornol. 59: catory devices for control of the face fly. I Econ. En/ornol. 726-32. 58: 39--41. Dorsey, C. K., H. E. Kidder & C. J. Cunningham. 1962. Face Hair, J. A. & E. C. Turner, Jr. 1965. Attempted propagation fly control studies in West Virginia in 1960 and 1961. I of Nasonia vitripennis on the face fly.I Econ. En/ornol. 58: &on. Entomol. 55: 369-74. 159-60. Drea, J. J. 1966. Studies of Aleochara tristis (Coleoptera: Sta- 1966. Susceptibility of mature and newly emerged face flies phylinidae), a natural enemy of the face fly. I Econ. Ento- to chemosterilization with apholate. I Econ. Entmno/. 59: mol. 59: 1368-73. 452-54. Eddy, G. W. & A. R. Roth. 1961. Toxicity to fly larvae of the Hammer, O. 1941. Biological and ecological investigations on feces of insecticide-fed cattle. I Econ. Entornol. 54: 408-11. flies associated with pasturing cattle and their excrement. Eltrigham, H. 1916. Some experiments on the house fly in Vidensk. Medd. Dan. Naturhist. Foren. Khobenhavn 105: 141- relation to the farm manure heap. I Agric. Sci. 6: 443-57. 393. (English) Engroff, B. W.,J. H. Knudsen & E.J. Hansens. 1972. Analysis Hansens, E. J. 1963. Face fly and its control. Leaflet 356. Ext. of the effects of some environmental factors on popula- Serv., Rutgers Univ., New Brunswick, New Jersey. tions of the face fly on dairy cattle. Environ. Entornol. I: Hansens, E. J. & P. Granett. ]963. Tests of Ciodrin and other 768-71. materials against face fly, Musca auturnnalis. I Econ. Ento- Fales, J. H. & O. F. Bodenstein. 1966. A promising new in- mol. 56: 24-29. secticide. Soap Chern. Spec. 42: 66-68, 104-06. Hansens, E. J. & I. Valiela. 1967. Activity of the face fly in Fales, J. H., O. F. Bodenstein & J. C. Keller. ]96Ia. Face fly New Jersey. I Econ. Entornol. 60: 26-28. laboratory rearing. Soap Chem. Spec. 37: 81-83. Hayes, C. G. & E. C. Turner, Jr. 1971. Field and laboratory 1961b. Face fly investigations in Maryland in 1961. Soap evaluation of parasitism of the face fly in Virginia. I Econ. Chem. Spec. 38: 85, 87, 89, 91, 109, 197. Entornol. 64: 443-48. Fales, J. H., O. F. Bodenstein, G. D. Mills, Jr & R. L. Urban. Heller, P. R. 1976. Biology of Aleochara tristis in relation to 1968. Face fly control up to date. Soap Chern. Spec. 44: 106, biological control of the face fly. Ph.D. thesis, Ohio State 108, 1I0, 112, 140--42. Univ., Columbus. 205 p. Fales, J. H., O. F. Bodenstein, J. D. Mills & J. H. Wessel. Holdsworth, R. P. 1962. Control of face flies attacking com- 1964. Preliminary studies on face fly dispersion. Ann. Ento- mercial dairy herds. J. Econ. Entornol. 55: 146--47. rnol. Soc. Arn. 57: 135-37. Houser, E. C. & C. W. Wingo. 1967. APhaereta palliPes as a Fales, J. H., O. F. Bodenstein, R. M. Waters & E. S. Fields. parasite of the face fly in Missouri, with notes on laboratory 1971. Insecticidal evaluations of d-trans-allethrin. Soap culture and biology. J. Econ. Entornol. 60: 731-33. Chern. Spec. 47: 64, 66, 68, 70, 135. Hower, A. A., Jr & T. H. Cheng. 1968. Inhibitive effect of Fales, J. H., J. C. Keller & O. F. Bodenstein. 1961c. Experi- Bacillus thuringiensis on the development of the face fly in ments on control of the face fly.I Econ. En/01ll01. 54: 1147- cow manure. J. Econ. Ento1ll01. 61: 26-31. 51. 1972. A field cage study of feeding behavior of the face fly Frishman, A. M. &J. G. Matthysse. 1966. Olfactory responses on dairy cattle. J. Econ. Entornol. 65: 602-04. 208 J. Med. Entomol. VoI.17,no.3

Jones, C. M. 1963. Research on the face fly during 1962. Proc. 1972. Boric acid effects on face fly fecundity. J. Econ. Ento- North Cent. Branch Entomol. Soc. Am. 18: 53. mol. 65: 740-41. 1967. Aleochara tristis, a natural enemy of face fly. L Intro- Lloyd, J. E. & J. G. Matthysse. 1970. Polyvinyl chloride-in- duction and laboratory rearing. J. Econ. Entomol. 60: 816- secticide pellets fed to cattle to control face fly larvae in 17. manure. J. Econ. Entomol. 63: 1271-81. 1969. Biology of the face fly: migration of larvae. J. Econ. Lodha, K. R., R. E. Treece & F. R. Kontz. 1970. Studies on Entolllol. 62: 255-56. the mat.ing behavior of t.he face fly. J. Econ. Entomol. 63: Jones, C. M. & J. G. Medley. 1963. Control of the face fly on 207-12. cattle with Co-Ral in grain and on pasture.}. Econ. Entomol. MacNay, C. G. 1961. Face fly, Musca autumnalis. Can. Insect 56: 214-15. Pest Rev. 39: 116,201,291. Jones, C. M. & J. M. Perdue. 1967. Heterotylenchus autumnalis, Matthew, D. L., R. C. Dobson & J. V. Osmun. 1960. The face a parasite of the face fly.}. Econ. Entomol. 60: 1393-95. fly is becoming a household pest. Pest Control 28: 16. Kantack, B. H., W. L. Berndt & E. U. Balsbaugh, Jr. 1967. Matthysse,1- G. 1961. Controlling the face fly on dairy cat.tle. Horn fly and face fly control on range cattle with aerial Stn. to Stn. Res. News 7: 1-2. applications of ultra-low-volume malathion sprays.}. Econ. 1962. The face fly, Musca autumnalis De Geer, on livestock. Entomol. 60: 1766-67. Chem. Spec. Manu! Assoc. Proc. Annu. Meet. 48: 149-53. Kaur, D. & P. C. Steve. 1969. Induced sterility in face fly- McGuire, J. U. & R. I. Sailer. 1962. A method of estimating face initial and sustained effects of hempa and metepa. J. Econ. fly populations on cattle. USDA, ARS-33-80. 8 p. Entomol. 62: 1462-64. Miller, 1- A., M. L. Beadles, J. S. Palmer & M. O. Pickens. Kaur, D. & B. C. Wentworth. 1972. Studies on the chemo- 1977. Met.hoprene for comrol of the horn fly: a sustained sterilization of the face fly. J. Econ. Entomol. 65: 21-23. release bolus formulation for cat.tle. J. Econ. Entomol. 70: Kearns, H. G. H. 1942. The control of flies in country and 589-91. town. Ann. Appl. Bioi. 29: 310-13. Miller, R. W. 1970. Larvicides for fly control-a review. Bull. Kessler, H. & E. U. Balsbaugh, Jr. 1972. Parasites and pred- Entomol. Soc. Am. 16: 154-58. ators of the face fly in east-central South Dakota.}. Econ. 1974. TH-6040 as a feed additive for control of the face fly Entolllol. 65: 1636-38. and house fly.J. £Can. Entomol. 67: 697. Kessler, H. & W. L. Berndt. 1971. Comparison of dust bags Miller, R. W. & L. G. Pickens. 1975. Feed additives for control to backrubbers for control of horn flies and face flies on of flies on dairy farms (Diptera: Muscidae). J. Med. Entolllol. beef cattle in east-central South Dakota. }. Econ. Entomol. 12: 141-42. 64: 1465-66. Miller, R. W., L. G. Pickens & L. M. Hunt. 1978. Methoprene: Killough, R. A. 1965. Effect of different levels of illumination field tested as a feed additive for control of face flies. }. on the life cycle of the face fly.}. £Can. Entomul. 58: 368- Econ. Entomol. 71: 274-78. 69. Miller, R. W. & E. C. Uebel. 1974. Juvenile hormone mimics Killough, R. A., J. G. Hartsock, W. W. & J. W. Smith. as feed additives for control of the face fly and house fly. 1965. Face fly dispersal, nocturnal resting places, and ac- J. Econ. Entornol. 67: 69-70. tivity during sunset as observed in 1963.}. Econ. Entomol. Miller, T. A. 1967. Some relationships of Musca autlllnnalis De 58: 711-15. Geer (Diptera: Muscidae): feeding, ovarian development, Killough, R. A. & E. C. McClellan. 1965. Face fly oviposition and incidence on dairy cattle. Diss. Abstr. B: 28: 2467. studies.}. Econ. Entmnol. 58: 716-19. Miller, T. A. & R. E. Treece. 1968. Some relat.ionships of face Knapp, F. W. 1962. Horn fly and face fly control studies with fly feeding, ovarian development, and incidence on dairy Dow M-1816.}. £Can. Entolllol. 55: 816-17. cattle. J. £Can. Ent01nol. 61: 251-57. 1965a. Free choice feeding of ronnel mineral block and Morgan, N. O. & L. G. Pickens. 1967. Cold tolerance ofadult.s granules for face fly, horn fly, and cattle grub control. .f. and pupae of the face fly. }. Econ. Entontol. 60: 1464-66. Econ. Entomul. 58: 836-38. Nickle, W. R. 1967. Heterotylenchus a.utumnalis sp. n. (Nema- 1965b. The effect of free-choice coumaphos salt mixtures on toda: Sphaerulariidae), a parasite of the face fly, Musca cattle and cattle parasites. }. Econ. Entomol. 58: 197-99. autulllnalis De Geer. J. Parasitol. 53: 398-40 I. 1966. Aerial application of trichlorfon for horn fly and face Ode, P. E. & J. G. Matthysse. 1964a. Face fly control experi- fly control on cattle. J. Econ. Entomol. 59: 468. ments.}. Econ. Entomol. 57: 631-36. 1967. Ultra-low-volume aerial application of trichlorfon for 1964b. Feed additive larviciding to control face fly.}. Econ. control of adult mosquitoes, face flies, and horn flies. }. Entontol. 59: 726-32. Econ. Entolllol. 60: 1193. 1967. Bionomics of the face fly. M'usca autwnnalis De Geer. Kobayashi, H. 1919. Flies in Korea. Report I. Chosen Igakukwai Cornell Univ. Mem. 402.90 p. Zasshi No. 24. 22 p. [Rev. Appl. Entulllol. Ser. B: 7: 142 Okamoto, H. 1924. The insect fauna of Quelpart. Island (1919).] (Saishiu-do). Bull. Agric. Exp. Stn. Gov. Gen. Chosen I. 2: 1- Krecob, M. A. 1949. Investigations on the biology of the nem- 4. atode, Thelazia rhodesi Demarest. Dohl. Akad. Nauk SSSR 66: Parrish, G. V. & R. R. Gerhardt. 1976. Daily face fly infes- 309-11. tation patterns on pasturecl beef cattle. Tenn. Farlll Home LaBrecque, G. C. 1961. Studies with three alkylating agents Sci. Prog. Rep. 97: 12-13. as house fly sterilants. J. Econ. Entolllol. 54: 684-89. Pechuman, L. L. & J. J. S. Burton. 1969. Seasonal distribution Lancaster, J. L., Jr, R. R. Blum & J. S. Simco. 1976. Labo- of Tabanidae (Diptera) at Texas Hollow, New York, in ratory evaluations of Onthophagus gazella against Musca au- 1968. Mosq. News 29: 216-20. twnnalis De Geer. Southwest. Entomol. 1: 111-13. Peterson, C., Jr & J. R. Boreherding. 1962. Face up to face Lancaster, J. L., Jr & J. S. Simco. 1975. Preliminary testing of flies and t.heir "stablemates." Successful Farming 60: 38-39. a synthetic pyrethroid against house and face flies. Arkansas Peterson, R. D. & H. J. Meyer. 1978. Trapping technique for Farlll Res. 24: 12. male face flies.}. Econ. Entol1lol. 71: 40-42. Lang, J. T. & R. E. Treece. 1971. Sterility and longevity effects Pickens, L. G. & A. B. DeMilo. 1977. Face fly: inhibition of of Sterculia foetido oil on the face fly. .J. Econ. Entomol. 64: hatch by diflubcnzuron and related analogues. J. Econ. 455-57. Entolllol. 70: 595-97. 1980 Pickens & Miller: Biology and control of the face fly 209

Pickens, L. G., R. W. Miller & J. J. Grasela. 1977. Sticky Smith, T. A., D. D. Linsdale & D. J. Burdick. 1966. An an- panels as traps for Musca autulllnalis. J. Econ. Entolllol. 70: notated bibliography of the face fly, Musca auturnnalis De 549-52. Geer, in North America. Calif Vector Views 13: 43-53. Pitts, C. W. & T. L. Hopkins. 1964. Toxico]ogical studies on Steve, P. C. & J. H. Lilly. 1965. Investigations on transmis- dichlorvos feed-additive formulations to control house flies sibility of Moraxella hovis in cattle by the face fly.j. £Con. and face flies in cattle feces. J. Eeon. Entolllol. 57: 881-84. Entomol. 58: 444-46. Poindexter, C. E. & T. R. Adkins, Jr. 1970. Control of the Stoffolano, J. G., Jr. 1968a. Distribution of the Het- face fly and the horn fly with self-applicatory dust bags. J. erotylenchus autumnalis, a parasite of the face fly, in New Eeon. Entolnol. 63: 946-48. England with notes on its origin. J. Eeon. Entomol. 61: 861- Poorbaugh, J. H., J. R. Anderson & J. F. Burger. ]968. The 63. insect inhabitants of undisturbed cattle droppings in north- 1968b. The effect of diapause and age on the tarsal accep- ern California. Calif. Vector Views 15: ]7-36. tance threshold of the fly, Musca autumnalis.J.lnsect Physiol. Roberts, J. E. 1963. Control of flies on beef and dairy cattle. 14: 1205-14. Ga. Agrie. Exp. Stn. Mimeogr. Ser. N.S. 186. II p. 1971. Be on the lookout for eyeworm infections. Dairy Digest 1965. Rubbing devices tJ'eated with insecticides for horn fly 37: 1,3. and face fly control on beef cattle. Ga. Agrie. Exp. Stn. Mi- Stoffolano, J. G., Jr & J. G. Matthysse. 1967. Influence of meogr. Sa. N.S. 227.9 p. photoperiod and temperature on diapause in the face fly, Roberts, J. E., P. E. White & D. M. Baird. 1963. Face fly and Musca autumnalis (Diptera: Muscidae). Ann. Entomol. Soc. horn fly population and control studies on cattle. Ga. Agrie. Am. 60: 1242-46. Exp. Stn. Mimeogr. Sa. N.S. 162. 13 p. Stoffolano, J. G., Jr & W. R. Nickle. 1966. Nematode parasite Robinson, J. V., H. W. Chambers & R. L. Combs, Jr. 1975. (Heterotylenchus sp.) of face fly in New York State. J. Eeon. Topical toxicity often insecticides to laboratory-reared Mus- Entomol. 59: 221. ca autulIlnalis. J. Econ. Entornol. 69: 89-90. Strickland, W. B., D. D. Linsdale & R. E. Doty. 1970. Hi- Rogoff, W. M. & A. L. Moxon. 1952. Cable type backrubbers bernation of face flies in buildings in Humboldt County, for horn fly control on cattle. J. Eeon. Entomol. 45: 329-34. California. Calif. Vector Views 17: 79-83. Ronald, N. C. & C. W. Wingo. 1973. Cost and effectiveness Teskey, H. J. 1960. A review of the life-history and habits of of horn fly and face fly control programs in semi-isolated Musca autumnalis De Geer. Can. Entornol. 92: 360-67. range herds in central Missouri. .J. Eeon. Entornol. 63: 693- 1969. On the behavior and ecology of the face fly. Can. Ento- 96. mol. 101: 56-76. Roussell, P. G. 1965. Comparative insecticidal susceptibility of Thimijan, R. W., L. G. Pickens & N. O. Morgan. 1973. Re- field-collected and laboratory-reared face flies, Musca au- sponses of the house fly, stable fly, and face fly to electro- IUlIlnalis. j. Econ. Entomol. 58: 674-76. magnetic radiant energy. J. Eeon. Entornol. 66: 1269-70. Rummel, R. W. 1971. Predation as a means of biological con- Thomas, G. D. & B. Puttler. 1970. Seasonal parasitism of the trol of certain coprophagous diptera. Diss. Abstr. B: 32: face fly by the nematode Heterotylenehus autumnalis in cen- 2779. tral Missouri, 1968. j. Eeon. Entomol. 63: ]922-23. Schmidtmann, E. T. 1977. Muscid fly predation by Vespula Thomas, G. D. & C. W. Wingo. 1968. Parasites of the face fly germaniea (Hymenoptera: Vespidae). Environ. Entomol. 6: and two other species of dung-inhabiting flies in Missouri. 107-08. j. Eeon. Entolllol. 61: 147-52. Schwarz, M., R. W. Miller, J. E. Wright, W. F. Chamberlain Treece, R. E. 1960. Distribution, life history, and control of & D. E. Hopkins. 1974. Compounds related to juvenile face fly in Ohio. Proe. North Cent. Branch Entolllol. Soc. Am. hormone. Exceptiona] activity of arylterpenoid compounds 15: 107-08. in four species of flies. j. Econ. Entornol. 67: 598-60 I. ]961. A comparison of the susceptibility of the face fly, Musea Seawright, J. A. & T. R. Adkins, Jr. 1968. Dust stations for autll1nnalis, and the house fly, M. domestica, to insecticides control of the face fly in South Carolina. j. Econ. Entomol. in the laboratory. j. Eeon. EnlO1nol. 54: 803-04. 61: 504-05. 1962a. DDVP applied regularly is helping control the face Shugart, J. I., J. B. Campbell, D. B. Hudson, C. M. Hibbs, R. fly. Ohio Farm Home Res., Ohio Agric. Ext. Stn. 47: 35,47. G. White & D. C. Clanton. 1979. Ability of the face fly to 1962b. Feed additives for control of face fly larvae in cattle cause damage to eyes of cattle. j. Eeon. Entomol. 72: 633- dung. j. Econ. Ent01nol. 55: 765-68. 35. 1964. Evaluation of some chemicals as feed additives to con- Singh, P., W. E. King & J. G. Rodriguez. 1966. Biological trol face fly larvae. j. Eeon. Entornol. 57: 962-63. control of muscids as influenced by host preference of 1966. Effect of bovine diet on face fly development. A pre- Macrocheles muscaedollles/icae (Acarina: Macrochelidae). j. liminary report. j. Eeon. Entmnol. 59: 153-56. Med. Entomol. 3: 78-81. Treece, R. E. & T. A. Miller. 1968. Observations on Hetero- Smith, T. A. & J. A. Krancher. 1974. Fifth supplement to an tylenchus autumnalis in relation 1:0 the face fly.j. Eeon. Ento- annotated bibliography of the face fly, Musca autumnalis mol. 61: 454-56. De Geer, in North America. Calif. Vector Views 21: 9-13. Turner, E. C., Jr. 1965. Area control of the face fly using self- Smith, T. A. & D. D. Linsdale. 1967. First supplement to an applicating devices.}. Eeon. Entomol. 58: 103-05. annotated bibliography of the face fly, Musca autumnalis Turner, E. C., Jr & J. A. Hair. 1966. Effect of temperature De Geer. in North America. Calif Vector Views 14: 74-76. on certain life stages of the face fly. }. Econ. Entomol. 59: 1968. Second supplement to an annotated bibliography of 1275-76. the face fly, Musca auturnnalis De Geer, in North America. ]967. Effect of diet on longevity and fecundity of laboratory- Calif. Vee/or Views 15: 119-21. reared face flies. j. Eeon. Entolnol. 60: 857-60. 1969. Third supplement to an annotated bibliography of the Turner, E. C., Jr & C. M. Wang. 1964. Residual and topical face fly, Musca autuml1alis De Geer, in North America. Calif· toxicity of certain insecticides to laboratory-reared face Vee/or Views 16: 119-23. flies.}. Econ. Entomol. 57: 716-19. 1971. Fourth supplement to an annotated bibliography of Uebel, E. C., P. E. Sonnet, R. W. Miller & M. Beroza. ]975. the face fly, Musca aU/llInnalis De Geer. Calif. Vector Views Sex pheromone of the face fly, Musca autll1nnalis De Geer. 18: 17-19. .J. Chem. Eeol. 195-202. 210 J. Med. Entomo!. Vol. ]7, no. 3

USDA. ]961. Face fly. Coop. Econ. Insect Rep. 11: 254. Weinmann, C. J., J. R. Anderson, P. Rubtzoff, G. Connally & ]963. Face fly. Coop. Econ. Insect Rep. 13: 870. W. M. Longhurst. 1974. Eyeworms and face flies in Cali- ]969. Face fly distribution. Coop. Econ. Insect Rep. 19: 728. fornia. Calif. Agric. 28: 4-5. ]975. Face fly distribution. Coop. Econ. Insect Rep. 25: 298. Wetzel, H. 1974. The significance of Musca autlllnnalis as a Vainshtein, B. A. & R. A. Rodova. ]940. Les ]ieux develop- disease carrier. Z. Parasitenkd. 39: 7]-72. ment des mouches de fumier dans les conditions dn. Tadzh. Wingo, C. W. ]970. Laboratory adaption of an indigenous Montagneux Med. Parasitol. 9: 364-68. [Rev. Appl. Entomol. braconid parasite of the face fly.J. Econ. Entomol. 63: 748- Ser. B: 31: 126 (1943).] 51. Valder, S. M., T. L. Hopkins & S. A. Va]der. ]969. Diapause Wingo, C. W., G. D. Thomas & N. M. Nelms. 1967. Labora- induction and changes in lipid composition in diapausing tory evaluation of two aleocharine parasites of the face fly. and reproducing face flies, Musca autumnalis. J. Insect Phys- J. Econ. Entornol. 60: ]514-17. iol. 15: ]]99-214. Wrich, M. L. 1970. Horn fly and face fly control on beef cattle Valiela, I. 1969. An experimental study of the mortality fac- using backrubbers and dust bags containing coumaphos or tors of larval Musca autumnalis De Geer. Ecol. Monogr. 39: fenthion. J. Econ. Entomol. 63: ] 123-28. 199-225. Wright, R. E. 1972. Nematode parasite of the face fly in On- Vockeroth, J. R. ]953. Musca autumnalis De Geer in North tario. Proc. Entomol. Soc. Onto 102: 168-75. America (Diptera: Muscidae). Can. Entomol. 85: 422-23. Wylie, H. G. 1973. Parasites of the face fly, Musca autumnalis Wallace, J. B. & E. C. Turner, Jr. 1962. Experiments for con- (Diptera: Muscidae), and associated Diptera near Belleville, trol of the face fly in Virginia. J. Econ. Entomol. 56: 214- Ontario. Can. Entomol. 105: 257-61. 15. Yendol, W. G. & E. M. Miller. 1967. Susceptibility of the face 1964. Low-level feeding of ronnel in a mineral salt mixture fly to commercial preparations of Bacillus thuringie1l.lis. j. for area control of the face fly, Musca autumnalis. J. Econ. Econ. Entomol. 60: 860-64. Entomol. 57: 264-67. Zapanta, H. M. & C. W. Wingo. 1968. Preliminary evaluation Wang, C. M. ]964. Laboratory observations on the life history of heliotrine as a sterility agent for face flies.J. Econ. Ento- and habits of the face fly, Musca autumnalis (Diptera: Mus- mol. 61: 330-31. cidae). Ann. Entomol. Soc. Am. 57: 563-69.