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Relative Effectiveness of Commmercial Fly Traps and Ammonium Carbonate/Yeast in Catching House Flies (Diptera: Muscidae)


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1 RELATIVE EFFECTIVENESS OF COMME RCIAL FLY TRAPS AND AMMONIUM CARBONATE/YEAST IN CATCHING H OUSE FLIES (DIPTERA: MUSCIDAE) By RYAN MERRILL WELCH A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2006

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2 Copyright 2006 by Ryan Merrill Welch

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3 ACKNOWLEDGMENTS I express deep gratitude to Dr Phil Koehler for his direction in this thesis and in my graduate study as a whole. His guidance a nd concern have been extraordinary. I also acknowledge my graduate committee—Dr. Chri s Geden, Dr. Phil Kaufmann, and Dr. Richard Patterson—for their help in making this thesis possible with revisi ons and general support. Also, thanks go to Dr. Phil Kaufmann for allowing space in his laboratory to rear house flies for my experiments. Ricky Vazquez, Jeff Hertz, and Matt Aubuchon offered valu able help in rearing house flies. Special thanks go to my friends and co-workers in the urban entomology lab, especially Gil Marshall and Osborne “Tiny” Willis, for their help in all things related to the urban laboratory. Debbie Hall was an invaluable help with so many aspects of my graduate experience from registering for classes to gene ral advice. Finally, I would like to thank my family for their lifelong support of my academic endeavors.

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4 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................3 LIST OF TABLES................................................................................................................. ..........6 LIST OF FIGURES................................................................................................................ .........7 ABSTRACT....................................................................................................................... ..............8 CHAPTER 1 REVIEW OF LITERATURE.................................................................................................10 Distribution and Classification...............................................................................................1 0 Morphology/Identification...................................................................................................... 10 Life Cycle..................................................................................................................... ..........10 Flight and Dispersal........................................................................................................... .....12 Mating Behavior................................................................................................................ .....13 Oviposition.................................................................................................................... .........15 Feeding........................................................................................................................ ...........16 Longevity...................................................................................................................... ..........17 Sex Ratio...................................................................................................................... ...........18 Economic Importance............................................................................................................ .18 Control........................................................................................................................ ............20 Fly Traps...................................................................................................................... ...........24 Objectives..................................................................................................................... ..........26 2 MATERIALS AND METHODS...........................................................................................27 Laboratory Rearing of the House Fly.....................................................................................27 Fly Trap Design Experiments.................................................................................................28 Scatter Experiment..........................................................................................................28 Commercial Fly Trap Experiment...................................................................................29 Fly traps....................................................................................................................29 Bioassay...................................................................................................................29 Attractant Experiments......................................................................................................... ..30 Aged Attractant Experiment............................................................................................30 Attractant Component Experiment..................................................................................31 Data Analysis.................................................................................................................. ........31 3 RESULTS...................................................................................................................... .........43 Fly Trap Design Experiments.................................................................................................43 Scatter Experiment..........................................................................................................43 Commercial Fly Trap Dimensions..................................................................................45

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5 Commercial Fly Trap Experiment...................................................................................46 Attractant Experiments......................................................................................................... ..47 Aged Attractant Experiment............................................................................................47 Attractant Component Experiment..................................................................................48 4 DISCUSSION AND CONCLUSIONS..................................................................................58 Fly Trap Design Experiments.................................................................................................58 Attractant Experiments......................................................................................................... ..61 Conclusions.................................................................................................................... .........64 LIST OF REFERENCES............................................................................................................. ..66 BIOGRAPHICAL SKETCH.........................................................................................................77

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6 LIST OF TABLES Table page 3-1 Percent escape of male, female, and tota l house flies into CDC ne t traps or laboratory room........................................................................................................................... ........50 3-2 Characteristics of six commercially availa ble fly traps used to evaluate house fly capture efficacy............................................................................................................... ...51 3-3 Percentage of male, female, and total house flies caught in six commercial traps............52 3-4 Percentage of male, female, and total house flies captured after 24 h in Trap n’ TossTM fly traps baited with an attractant aged up to 4 d...................................................53 3-5 Percentage of house flies caught in baited Trap n’ TossTM fly traps with attractants aged up to 4 d................................................................................................................. ....54

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7 LIST OF FIGURES Figure page 2-1 CDC Net Trap used to capture hous e flies escaping from screen cages............................32 2-2 Bottom-entry release cage design fitted with a Trap n’ TossTM cone and open to laboratory room................................................................................................................ ..33 2-3 Top-entry release cage design fitt ed with modified Trap n’ TossTM and open to laboratory room................................................................................................................ ..34 2-4 Trap n’ TossTM fly trap.....................................................................................................35 2-5 The Advantage Fly TrapTM................................................................................................36 2-6 BC 1752 Dome (McPhail) Trap.........................................................................................37 2-7 Rescue! Reusable Fly Trap.............................................................................................38 2-8 Victor Fly Magnet Trap..................................................................................................39 2-9 Fly Terminator Pro........................................................................................................ ..40 2-10 Bottom-entry study design with Trap n’ TossTM...............................................................41 2-11 Top-entry study design with Victor Fly Magnet Trap....................................................42 3-1 Effect of attractant age on percent cat ch of male, female, and total house flies................57

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8 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science RELATIVE EFFECTIVENESS OF COMME RCIAL FLY TRAPS AND AMMONIUM CARBONATE/YEAST IN CATCHING H OUSE FLIES (DIPTERA: MUSCIDAE) By Ryan Merrill Welch December 2006 Chair: P.G. Koehler Major Department: Entomology and Nematology House flies are important pests that are best controlled by using integrated pest management (IPM) because of the house fly’s abili ty to develop insectic ide resistance and its considerable reproduction poten tial. Commonly used IPM tactics include sanitation, insecticides, biological contro l agents, and fly traps. My studies deal with the ab ility of commercial fly trap s to catch house flies and the evaluation of house fly behavior in relation to fly trap catches. In a series of experiments, house flies were released into screen ed cages fitted with commercial fly trap funnels. The released house flies escaped the cage either into the laboratory room or into fly traps. However, despite the seemingly random escape from the release ca ges, air movement may also affect house fly dispersal behavior. In my expe riments, more house flies scattere d into the laboratory room than were captured in the un-baited CDC net trap, and more house flies were captured in the un-baited CDC net trap than in the un-baited Trap n’ TossTM. The behavior of th e house flies in my experiments showed the house flies’ tendency to fly upwind. Six commer cial fly traps were compared for their ability to cap ture house flies released into sc reened cages. The Trap n’ TossTM fly trap caught significantly more house flies than the Fly Terminator Pro fly trap, and this was probably due to a number of design f actors including a large entrance hole area, a small

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9 exit hole area, a large ratio of entrance to exit hole, the cone slope being close to 60o, and the color of the entrance cone. Two commonly used materials in commercial fly trap attractants, yeast and ammonium carbonate, were tested to identify th eir attractancy to adul t house flies released into screen cages. In my studies, yeast was more attractive than water at 0, 2, and 4 days old by ~30-35%. My studies also found ammonium carbonate was never more attractive than water, and when added to yeast did not increase attractancy. Theref ore, ammonium carbonate was not a chemical attractant for the adult house fly when used alone or in conjunction with the other attractants tested. Yeast, being a living or ganism, had a life cycle that wa s subject to availability of nutrition. The attractant combination (yeast a nd ammonium carbonate in a 41.67:1 ratio) in my study increased linearly from day 1 to 4. My resu lts suggest that when using yeast to attract house flies, as long as the yeast is protected from temperature extremes, house fly attraction will increase linearly for at least 4 days. My studies have shown that commercial fly traps have gr eat potential as house fly IPM tools. They are versatile devices to be used in IPM programs, and can be used both as control devices and for monitoring house fly populations. According to my laboratory studies, and with the further exploration with fiel d studies, commercial fly trap s represent a viable tool for managing house flies in a wide variety of sett ings, including agriculture, urban, and military.

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10 CHAPTER 1 REVIEW OF LITERATURE Distribution and Classification The house fly, Musca domestica L., belongs to the kingdom Animalia, phylum Arthropoda, class Insecta, order Diptera, suborder Cyclorrhapha, family Muscidae, subfamily Muscinae (West 1951). Linnaeus described the sp ecies in 1758. Other common names include typhoid fly, cholera fly, dysentery fly, and enteric fly after important diseases thought to be vectored by the fly at certai n times and places (West 1951). The house fly is for all practical purposes considered worldwide in its distribution, although it may not be found in arctic regions or places of high altitude (West 1951). The fact remains that it is a nearly ubiquitous, synanthropic insect that is found fr om sub-polar regions to the tropics (Graham-Smith 1914, West 1951). Morphology/Identification The house fly is a gray insect that is usually 4 to 9 mm long depending on available resources. The wings are held di rected posteriorly when at rest. There is a sharp bend in the fourth longitudinal wing vein. Th ere are four darkened, parallel stripes running vertically down the thorax (West 1951). The abdomen is a gray to yellow color with a dark midline and dark markings set irregularly. Females are usually slightly larger than males and are easily distinguished by noting the space between the eyes, in which the females’ is almost twice as large (West 1951). Life Cycle The house fly, like all Diptera, is holometabolous with distinct egg, larva, pupa, and adult stages (Chapman 1998). The first stage in any insect development, including M. domestica is the egg. The eggs are white a nd appear polished, due to a cl ear, viscous coating (Newstead

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11 1908, West 1951). They are about one millimeter in length and have a pattern of small hexagonal markings due to the shape of the egg tube in which the ovum developed (West 1951). The egg may hatch in as little as 7 hours and 35 minutes when the temperature is an optimum 34 to 38oC and in as long as three days if the temperature is as low as 10oC (Hewitt 1914, Larsen and Thomsen 1940). The usual time to hatch is be tween eight and 20 hours. Eggs that become too dry will not hatch. Larvae, or maggots, emerge from the egg by way of a slit that appears on the dorsal side of the egg (West 1951). The larvae are white to cr eam-colored and possess no eyes, legs, antennae, or other appendages (West 1951). The larva mu st pass through three instars reaching 7-12 mm before pupating (Newstead 1908, West 1951). The thir d instar migrates from the food source to a dry, cool place to pupate. The larval stage can ta ke anywhere from five days to eight weeks, depending on the temperature and food quality (Newstead 1908). The fly reaches the pupal stage when the moveme nts of the contracted larvae have ceased and it is still a cream color, and ends when the adult fly emerges (Larsen and Thomsen 1940). The puparium color quickly changes from the crea m color darkening to a reddish brown or black (Newstead 1908, West 1951). It is usually ar ound 6 to 7 mm long and weighs about 21 mg (Newstead 1908, West 1951). The pupal stage typi cally accounts for more than half of the immature development time of the fly. It ma y last anywhere from 3.75 to 28 days, depending on the food of the larval stage and the temperature (Newstead 1908, Larsen and Thomsen 1940). Development time is inversely proportionate to temperature to a certain threshold. The upper temperature range in which retardation becomes apparent is between 36 and 40oC (Larsen and Thomsen 1940). If the medium reaches temperature below 10oC, there is a considerable level of mortality (Sacca 1963).

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12 The adult, or imagio, emerges from the puparium by breaking away the anterior end with the ptilinum found on the frontal region of th e head (Newstead 1908, West 1951). The fly escapes the puparium and escapes the pupal medi um such as manure, debris, sand, etc. by contracting the ptilinum (West 1951). This impo rtant structure is expanded and contracted by changes in blood pressure and retrac tion of muscles (Laing 1935, West 1951). The newly emerged adult crawls about rapidl y as the wings unfold and the exoskeleton becomes hard. The wings become flat, thin, and transparent by withdrawal of fluids and are supported by the veins. At last the ptilinum is withdrawn an d only the frontal “lunule” that marks its former location is left behind (New stead 1908, West 1951). Overall, the whole process from egg to adult, like every life stage, varies with temperature. However, when the medium is 33.22oC, the process takes about 6.5 days (Hogsette 1995). Flight and Dispersal House flies are able to fly considerable distance s. The extent of this distance is different according to different authors. Bishopp and L aake (1921) found that within 24 hours of release, house flies average a distance of about 9.66 km, with a maximum distance of about 21.15 km. Schoof and Siverly (1954) conducte d a study that suggests that flie s can travel as far as 32.19 km by way of flight. They also found that large numbers of flies might migrate up to 6.44 km. However, most flies are not reco vered at distances greater than 3.22 km. This is because house flies have been shown to fly in a more or less random pattern, until a suitable place is located (Sacca 1963). The house fly tends to travel shorte r distances in urban areas where structures, sources of food, and sources of oviposition are more readily available, whereas in rural environments, the fly will have to disperse fu rther to find suitable sites (Hindle 1914, Murvosh and Thaggard 1966). Additionally, air temperature limits the flight ability of the house fly. As temperature increases, so does th e motility of the house fly (Sacca 1963).

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13 Arguably the most important means of dispersa l for the house fly, and almost certainly the reason for its worldwide distributi on, is hitchhiking. Flies can tr avel great distances by using large mammals or human-assisted transportation (Sacca 1963). Mating Behavior The mating behavior of the house fly is a comp licated process that, once sexual maturity is reached, is always initiated by the male. Many researchers have investigated this issue. However, the temperatures used in their studies we re not always the same. Males require at least 16 hours to mature, whereas females require at least 24 hours (Murvosh et al.. 1964a). Chang (1965) determined a male maturation time of at least 20 hours and female maturation time of at least 40 hours. Despite the fact that female flie s may not be ready to copulate, males persist. This may lead to a female fending off and possi ble damaging of the male, especially his wings, with her posterior legs that are held near the tip of the abdomen (Bishopp et al. 1915). Copulation may occur much later even though males aggressively try. Bishopp et al. (1915) found flies mating as late as 16 days after emergence. This large range of time may be, at least in part, the result of nutrition. The nutrition may affect either the sexual development or the desire to mate. Bishopp et al. (1915) observed mating within four days of emergence on flies fed with milk and horse manure, while a group fe d partially ripened peaches and horse manure required 16 days. Copulation was no t observed among the unfed flies. Sexually determined males seek females us ing visual cues. The male is sexually aggressive, and will attempt to mate with anyt hing resembling a female house fly, including male house flies, fly cadavers, and in animate objects (Rogoff et al. 1963). However, visual cues are not necessary for a male to locate a mate. In fact, mating can occur in complete darkness (Basden 1947, Rogoff 1965).

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14 The male fly also uses chemical cues to id entify a partner (Patterson 1957). Both male and female house flies have gender-specific hydrocar bons, as well as hydrocarbons common to both sexes (Nelson et al. 1981). The major pheromone of the house fly is found on the female. It is a C23 hydrocarbon called (Z)-9-tricosene (Carlson et al. 1971). The ac tivity of this pheromone is enhanced by other hydrocarbons including methyl alkanes, (Z)-9,10-epoxytricosane, and (Z)-14tricosen-10-one (Uebel et al. 1976, Nelson et al. 1981, Bloomquist et al. 1984). It works as a short-range attractant, sex-rec ognition factor, arrestant, and copulatory stimulant of males (Mayer and James 1971, Mans ingh et al. 1972, Carlson et al. 1974, Adams and Holt 1987, Hanley et al. 2004). When the male does locate an appropriate partner, both flies begin seemingly random movements. The male will stop movement when he gets within a few centimeters of the female, wherein he leaps on her back (Rogoff et al. 1963). The mount may also be attempted in the air, but the couple quickly comes to a rest, as copu lation never happens in flight. If disturbed, however, the couple may fly to another surface toge ther (Murvosh et al. 1964a). Mating occurs in the superimposed position, with the male above the female and both facing the same direction. If the male is too small or disturbed, it may face at a near 90o angle or even face the opposite direction (Hardy 1944, West 1951). After mounting, the male caress es the head of the female with the tarsi of his front legs This probably has a reflex e ffect on the female as it usually results in ovipositor extension unt il it contacts the male genital atrium. The aedeagus extends as does the accessory copulatory vesicles of the female so that they contact each other. This is when the spermatozoa make their way into the spermethecae of the female (West 1951). The spermatozoa are transferred to the female in as little as three to five minutes however, the couple will usually stay in coitus fo r a much longer time (Murvosh et al. 1964a). Coitus has been

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15 observed to last anywhere fr om 30 minutes to two hours (H ampton 1952, Murvosh et al. 1964a, Chang 1965). After mating, the female usually remains mono gamous, and only a few mate more than a few times (Riemann et al. 1967). In a study by Zi mgrone et al. (1959), only two percent of the females mated multiple times. One fertilization episode is sufficient for the entire egg-laying period (Zimgrone et al. 1959). The male seminal fl uid, and not mechanical stimulation or sperm, has been shown to cause the loss of fema le sexual receptivity (Riemann et al. 1967). Oviposition Many factors are involved in ovigenesis and ovi position. For instance, diet is of utmost importance. Protein is necessary for the production of mature e ggs in the female house fly, as with many other dipterans (Sacca 1964). Whereas the diet is integral in the production of eggs, it is also important insofar as how quickly or slowly the eggs ar e oviposited. Bishopp et al.. (1915) provided one group of flies peaches for food as oppos ed to another lot bei ng supplied with milk, peaches, and horse manure. Those fed peaches re quired 16 days to oviposit, whilst those fed the mixture of foods took five. This suggests th at variety, or probably more specifically, the presence of protein, in the diet al.lows the female to oviposit more quickly. Temperature also plays an important role in the rapidity of egg deposition. In general, within reasonable ranges, speed of egg deposit ion is positively correlated with temperature (Bishopp et al.. 1915). Oviposition w ill altogether cease if the atmo spheric temperature drops to 10o C or lower, regardless of the substratum temperature (Kalandadze and Chilingarova 1942, West 1951). Humidity is also a factor in time to first ovi position. As observed w ith temperature, when humidity increases, the speed of egg deposition increases. The humidity may make the media is more moist, and therefore more appealing to the female flies (Bishopp et al.. 1915). However,

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16 because liquid medium is hardly ever a suitable location, there appears to be an upper limit to this effect (Kalandadze and Chilingarova 1924). Taking into account the variety of factors impacting egg deposition, a female house fly may require from five to twenty days post-eclosi on to deposit her first batch of eggs (Griffith 1908, Bishopp et al.. 1915). Once she is ready to ovi posit, she must find an appropriate medium. An appropriate medium is one that will provide am ple nutrition to the larvae that will hatch from the eggs. The larvae develop most quickly in any site that is fermenting and vegetal in nature. Flies often oviposit in garbage and manure (Mellor, J.E.M. 1919, Kalandadze and Chilingarova 1942, West 1951, Sacca 1963, Cosse and Baker 1996). Another factor influencing oviposition is the presence of other flies. Jiang et al. (2002) documented increased oviposition at sites where other female flies were present, possibly due to the semiochemicals n-tricosane and (Z)-9tricosene. When a suitable spot is located, a fe male fly will orient to cracks and crevices in where she either inserts her oviposit or, or crawls into the crevice so that the eggs are protected from desiccation and actinic light (Newstead 1908, West 1951). House fly reproductive capacity is considerable Hodge (1911) calculate d that if one pair of flies, unchecked, began reproducing in April, by August there would be 191,010,000,000,000,000,000 flies. It is estimated that this would cover the surface of the earth to a depth of 47 feet. One female fly usually ov iposits anywhere from 75 to 159 eggs in batches (Newstead 1908, Dunn 1923, Herms and James 1961). House flies have been reported to produce between two and twenty-one egg batches at intervals of eight to fourteen days, though usually only two to four ba tches are deposited (Griffith 1908, Bishopp et al. 1915, Dunn 1923). Feeding House flies, like all an imals, require nutrition in order to maintain body processes, such as flight, movement, feeding, etc. a nd to provide for the building bloc ks of the organs, especially

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17 the reproductive system. For the house fly, suga rs or assimilable star ch is required, while proteins or byproducts of protein hydrolysis are necessary for organ formation (Kobayashi 1934, West 1951). Certain substances and chemicals are f eeding attractants. Although house flies are omnivorous, there are suitable, but unattractive, foods that require stim ulus for consumption. The sugars that are particularly attractive to a dult flies for feeding purpo ses include lactose and dextrin. Sugars that are not n ecessarily attractive despite their nutritive potential are glucose, maltose, starch, or sucrose (Richardson 1917). Richardson (1917) also found that succinic and lactic acid were feeding stimulants. Awa ti and Swaminath (1920) found ammonia, hydrogen sulfide, and phosphorous compounds to be adequate to induce feeding in the house fly adult. Robbins et al. (1965) found casein, more specific ally, its amino acid components (leucine, lysine, isoleucine, and methionine) and the guanonsine monophosphate of yeast hydroloyzate to be feeding stimulants. Regardless of how attractive or nutritive a food substance is to the house fly, it can only be consumed if it is in a liquid or semi-liquid form, or is capable of being rendered so. Some solid foods are, if soluble, put into solution by the house fly’s use of vomit from the crop. There are some instances where non-soluble solids will be taken up, then broken down further with prestomal teeth. After the food is ingested, it will make its wa y through the alimentary tract (Sacca 1963). Longevity The longevity of house flies, no doubt, is direc tly related to the temp erature and humidity. West (1951) states that in the summer flies live no longer than a few weeks, whilst in the colder months, three months is realistic. Many studi es have examined house fly longevity. Herms (1928) recorded a mean of 30 days. Murvosh et al. (1964b), found that at a mean of 23.9o C,

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18 males live an average of 33 days, and females 43 days, with a range of one to 99 days. Rockstein (1957) found that wh en the temperature was 26.7oC and relative humidity was 45%, male house flies lived from 15 to 40 days, and female house flies lived from 20 to 60 days. The age of the parents affects the longevity of the offspring. The average life span of the succeeding generations decreases if only eggs fro m young flies are used. Flies developing from females aged 20 to 30 days at oviposition resu lted in an increase in the average lifespan (Callahan 1962). Sex Ratio Studies with lab colonies suggest that the sex ratio of house f lies is about 1:1. Murvosh et al. (1964a) reported a sex ratio of 50.6 to 49.4 rati o based on 8700 individuals, and a ratio of 53.5 to 46.5, based on 5233 individuals. Rogoff (1965) de termined his sex ratio using 16 different samples of 100 pupae each, and reported similar results. West (1951) observed that when the average a dult size is smaller than 6-7 mm in length, then males would predominate, while when the average adult size is larger, the males never predominate the population. Herms (1928) showed, while working with Lucilia sericata (Meigen), that larval females re quire more nourishment than their male counterparts to survive to the pupal stage. This being the cas e, an abundance of males may signi fy a lack of larval nutrition in the area. Economic Importance House flies gather at garbage, compost, sewage, manure, livestock facilities, and other unsanitary sites, and then readil y enter buildings making them a considerable nuisance to people and animals (Chapman et al. 1998). Although the house fly is not as much of a problem in many advanced industrial societies due to personal transportation being automobiles instead of horses, it is still one of, if not the most, important pests in markets, stores, and animal sites due mostly to

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19 annoyance (Cosse and Baker 1996, Robinson 1996, Chap man et al. 1998, Hogsette 2003). They become most annoying when they land on people and animals causing irritation. When they crawl on food the potential for pa thogen transmission is raised Whilst resting on walls and windows they leave behind specks from vomit a nd feces. Mere annoyance has turned into litigation. When flies from lives tock facilities find their way in to the spreading suburban and urban areas, lawsuits follow (Campbell 1993). Th e annoyance that the flies cause livestock may also reduce production yields (Cosse and Baker 1996, Chapman et al. 1998). House flies are economically important as dis ease vectors. House flies have been long suspected of disease transmission (Howard 1911). However, it was not until the studies carried out by Watt and Lindsay (1948) a nd Lindsay et al. (1953), that there was convincing evidence for the house fly’s involvement. After a fly has become infected with a pathogen, it can transmit the disease one of four ways. These include co ntact with the body set ae, setae of the feet, regurgitation, and passage through the digestive tract (West 1951) While West (1951) and Greenberg (1973) indicate that of these four, th e passage through the dige stive tract is probably the most important because it allows for multip lication of the disease organism within the alimentary canal, Zurek (2001) shows that the po tential for a pathogen to be spread via vomitus or feces is dependent on the specific pathogen. Greenberg (1971) lists over 30 viruses, hundreds of bacteria species, as well as a number of fungi, protozoa, and nematodes that are associated with M. domestica A far from exhaustive list of some of the disease organisms that house flies are known to transmit are Escherichia coli (including O157:H7), Shigella spp., Salmonella spp., Helicobacter pylori, Yersinia pseudotuberculosis and the organisms responsible for typh oid fever, cholera, summer diarrhea,

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20 dysentery, tuberculosis, anthrax, opthalmia, and parasitic worms (West 1951, Greenberg 1971, Grubel et al. 1997, Iwasa et al. 1999, Zurek et al. 2000, Geden 2005). Control The management of the house fly is an exercise in problem solving. West (1951) broke the control of house flies into two cat egories: planned control and emer gency control. This approach is better known as integrated pest management (IP M) today. West’s appare nt foresight into pest management is due to the fact that managing house flies offers a classic opportunity for IPM. An effective house fly management strategy incl udes sanitation and ot her cultural methods, mechanical control, insectic idal control, and, more recen tly, biological control (West 1951, Kolbe 2004). Greene and Breisch (2002) documented the success of IPM in urban environments by comparing the number of pest control service requests. From 1988 to 1999, the requests for the group that included house flies we re decreased to half its orig inal number. Also, the amount of insecticide used was drastically decreas ed to about 5% of its original use. Sanitation is the primary m eans of managing house fly pr oblems by eliminating breeding sites and larval food, which also attract adult flies (Clymer 1973, Gojmerac 1979, Meyer and Shultz 1990). Thus, by eliminating trash and manure, all stages of the fly life cycle are disrupted. In the urban setting, the main breeding site for the house fly is garbage. Kolbe (2004) offers insight into the management of garbag e for house fly control. Food in trash cans and dumpsters should be picked up frequently. Twic e a week is ideal. Trash cans and dumpsters should have tight-fitting lids, and should be kept closed. The ga rbage placed in trash cans and dumpsters should be sealed off in plastic bags to minimize attractive odors. Dumpsters should be kept as far away from building entrances as possible. Also, keeping the dumpster and the ground underneath it clean will help reduce house fly numbers.

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21 In the agriculture, the main br eeding site is manure (West 1951). Agricultural settings that have accumulated manure have higher house fly populations (Lysyk and Axtell 1986). Feedlots should allow for proper drainage that minimizes we t spots and prevents the build up of waste. Manure and spilled feed should be removed twice a week (Clyme r 1973). Sanitation, especially the removal of manure, of poultry houses and dairies is also an essential tool in an integrated pest management strategy against house flies (Lysyk and Axtell 1986, Lazarus et al. 1989). Manure removal is most effective in all instances when it is done in the late spri ng (Axtell 1970a). Also the bedding material that is r outinely excreted upon by livestock may be changed or added tosawdust or sand-in order to make the manure le ss enticing as a breedi ng site for house flies (MacCreary and Haenlein 1962, Schmidtmann 1988, Lazarus et al. 1989). Another cultural method to prevent house fly problems is exclusion. House flies are anthropomorphic and move to human structures through temperature and odor attractants. A way to combat this invasion is to make sure th at all windows and doors have screening, that the screens lack holes, and are prope rly fitted (West 1951). House f lies can also be excluded by fans and air doors between rooms (West 1951, Hedges 1994). Mechanical control is anothe r method critical to an e ffective house fly management regimen. Most likely the oldest control meas ure for house flies falls under this category flyswatting (West 1951). More modern methods of mechanical control are the plethora of available fly traps. The different types of fly traps include baited traps, light traps, and sticky traps (West 1951, Pickens et al. 1975, Skovma nd and Mourier 1986, Pickens 1989, Kaufman et al. 2005b). Of these types of traps there are a vari ety of different designs, attractants, and killing methods. These are effective supplements to other control methods because they are able to trap flies up to 24 hours a day, seven days a week with minimal effort (Kolbe 2004).

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22 Insecticides can be used to tr eat a house fly problem. Chemicals have been used to control flies since before World War II (West 1951). But, this should not be done indiscriminately, but rather after other methods have failed to reduc e the fly populations (He dges 1993). This is due to the house fly’s ability to develop resistance to almost every insecticide used against it (Learmount et al. 2002, Scott et al. 2000, Gao and Scott 2006). Not only does overuse of insecticides contribute to house fly resistance, but there are also the concerns of public perception, increased cost, safety, and envir onmental considerations (Carson 1962, Appling 1988, Stinson 1989). Different formulations should be considered for house fly management based on the situation. They are residual insectic ides applied to adult resting sites and larval breeding sites, baits, and aerosols and ultralow-volume (ULV) applications used as space treatments (Kolbe 2004). Insect icide treatment should be used with caution as it may hinder a proper IPM program i.e. affect natural predators th at could be used as bi ological control agents (Wills et al. 1990). Augmentative biological control is the newest form of house fly control and can be used as part of a successful IPM program. Fly egg predat ors represent one type of biological control, the two major species being Carcinops pumilio (Erichson) and Machrocheles muscadomesticae (Scopoli) (Axtell 1970b, Geden and Stoffolano 1987). Carcinops pumilio is capable of causing ~95% mortality of house flies, whereas M. muscadomesticae can cause ~80% mortality in the lab, with somewhat lower results in the field due to abiotic factors and th e presence of alternate prey (Geden and Axtell 1988, Geden et al. 1988). Fly larval pred ators also exist as biological control agents. The black dump fly, Ophyra aenescens (Wiedemann), is a predator of house fly larvae and is capable of out-competing the house fly (Nolan and Kissam 1985). Although O. aenescens is a filth fly, it remains a suitable candidate for biological control because it is not a

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23 nuisance. This is due to the adult habit of staying close to the breeding medium and its preference for woods and shade as opposed to human structures (L egner and Dietrick 1974, Nolan and Kissam 1987). Muscovy ducks are predator s of adult house flies that may be used as a biocontrol agent. The ducks ingest about 25 ho use flies per 15 minutes, and controlled 90% of the house flies in a much shorter time than fly traps or baits (Glofcheskie and Surgeoner 1990). Another group of biological cont rol agents are the Pteromalid parasitoids that attack house fly pupae. The most common are Muscidifurax raptor Girault and Sanders, Muscidifurax zaraptor Kogan and Legner, Muscidifurax raptorellus Kogan and Legner, Spalangia cameroni Perkins, Spalangia endius Walker, and Spalangia nigroaenea Curtis (Rueda and Axtell 1985, Havron and Margalit 1991, Geden and Hogsette 2006) These are naturally occurring species that hardly ever reach peak numbers and activity before or during the peak house fly nuisance levels, therefore, the parasitoid populations must be augmented either by inoculative or inundative release (Legner and Brydon 1966, S kovard and Nachman 2004). Skovard and Nachman (2004) reported that the inundative release of S. cameroni usually caused house fly populations to remain under nuisance levels (1325 house flies per animal) on both dairy and pig farms. Single species releases of Muscidifurax spp. and Spalangia spp have had mixed results (Geden et al. 1992, Geden and Hogsette 2006). Combined releases of the two genera are preferred due to Muscidifurax spp searching for hosts on the surface of manure, and Spalangia spp searching deeper (Rueda and Axtell 1985). Muscidifurax raptorellus and S. cameroni simultaneously released accounted for up to ~80% pupal mortality (Geden and Hogsette 2006). Microbes can also provide biological cont rol of the house fly. One example is the entomopathogenic fungus, Entomophthora muscae Cohn. The fungus causes adult house fly death in about 5-8 days after infection (Zurek et al. 2002). Once inf ected, house flies have a

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24 100% mortality rate (Steinkraus and Kramer 1987). Infection of othe r individuals is likely due to the males’ preference to mate with infected fe males (Zurek et al. 2002). Another aspect that makes E. muscae a good biological control ag ent is its ability to disc harge conidiophores from the cadaver of the dead fly (Zurek et al. 2002) A second entomopathogenic fungus that offers promise is Beauveria bassiana (Balsamo). In laboratory studie s, up to 98% mortality of house flies after 15 days of exposure has been reported (Lecuona et al. 2005). In the field, B. bassiana treated facilities had lower house fl y populations than facilities tr eated with pyrethrins (Kaufman et al. 2005a). Nematodes in the genus Heterorhabditis Steinernema and Paraiotonchium have been tested against house flies in the laboratory. While some show the ability to infect house flies and show promise as control agents in the lab ( Paraiotonchium muscadomesticae Coler caused 98% mortality with a huge concentration of nematodes) the nematodes’ infectivity is highly reduced in manure, requiring larger application con centrations (Mullens et al. 1987, Geden 1997). Fly Traps There are three common types of house fly traps: light traps, sticky traps, and baited traps. Light traps consist of an ultraviolet lamp that attracts the house fly into an electrocuting grid (ELT) or a glue board (GLT) (Pickens 1994) Although ELTs are successful, GLTs are becoming increasingly popular due to the noise emitted every time an insect is killed, and because when electrocuted, house flies can sc atter body parts and bact eria throughout the air (Pickens 1989). Though Kolbe (2004) reports that low-voltage ELTs do not introduce fly fragments into the air, Pickens (1989) observed a s catter of parts up to 1.5 m from the trap. Light traps should be placed 0.3 to 0.9 m from the ground about 15 m apart around common fly gathering places to maximize their effectiven ess (Driggers 1971, Gilbert 1984, Pickens 1994). Pickens et al. (1969) reported almost 100% capture of hous e flies in a 7.6 m2 room in only seven

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25 hours. However, Thimijan et al. (1970) reported 0 to 16.3% catch of released house flies. It is not surprising that some authors suggest that light traps are not sufficient control devices and should only be used for monitoring (Schie fer 1970, Skovand and Mourier 1986, Rambo 1995). Another type of trap that may be employed in an IPM program for house fly management is sticky traps. One common type of sticky trap is a roll of paper with glue adhesive, usually made attractive by color or prin ting patterns such as other flie s on it. Kaufman et al. (2005b) found that these traps caught over 80,000 house flies on each of five dairy calf greenhouse facilities in only four days. They concluded that the sticky trap was responsible for a decline in house fly populations and in each case, the prod ucer noticed a reduction in house fly numbers and was pleased. However, the efficacy of this type of trap declined in dusty environments, such as occurs on many poultry farms (Kaufman et al. 2001). Another common type of sticky trap is a pyramid sticky trap. Given the numbers of hous e flies trapped, these se em to be better suited for monitoring instead of control. Pickens a nd Miller (1987) caught 5.6 to 14.0% of released house flies with pyramid traps. They also point out that after about two days, the adhesive became saturated with flies and was unable to cat ch significant numbers. Pickens et al. (1986) and Johnson and Campbell (1987) de scribe pyramid traps as being most useful as monitoring devices, not control devices, for Musca vetripennis (Say) and Musca autumnalis De Geer respectively. Sticky tapes are also used as m onitoring devices (Pickens et al. 1972, Turner Jr. and Ruszler 1989). Baited traps are probably the ol dest of the fly traps, having been in use since about 1911 (Pickens 1995). The baited fly trap consists of an odiferous attractant in side of the trap that entices the house flies to enter through an inverted cone (Bishopp and Henderson 1946, Pickens 1995). Similar to the other trap types, there is debate whether baited tr aps should be used as

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26 monitoring or control devices. Dadour and Cook (1992) point out that to reduce a fly population by 50-90%, a trapping regimen must remove 24-58% of the house flies. Pickens and Miller (1987) report catching only 4.4 to 20.0% of releas ed flies in baited trap s. Smallegange (2004) states that although baited traps show promis e, they are not yet effective enough to reduce populations to acceptable levels. Das (1994) report ed that using Redtop Flycatcher baited with ready made protein meal was easier and less expensive to use than a Baygon bait, but admitted that it did not catch as many hous e flies as the Baygon bait kille d. Burg and Axtell (1984) and Geden (2005) also refer to ba ited traps as monitoring devices. Despite Dadour and Cook’s (1992) pessimism, Cohen et al. (1991) atta ined 64% reduction in house fly density, and attributed a 42% reduction in dia rrhea visits and 85% reduction in shigellosis visits of Israeli soldiers to the trapping. Perhap s some of the low baited trapping numbers can be attributed to how the baited traps are pl aced, after all, daily catches of up 40,000 to 129,000 have been reported (Blair 1945, Pickens 1995). Also, season al catches of 1.1 to 9.7 million have been recorded (Fenton and Bieberdorf 1936, Mer a nd Paz 1960). Baited traps should be placed outside every 6-9 m and placed as close as possi ble to common fly-breeding materials (Pickens 1994). Traps are most effective when placed within 0.3 m of the ground (Mitchell and Tingle 1975). Objectives The first objective of these experiments was to discover the relative effectiveness of each of six commercial fly trap de signs in capturing house flies. The second objective was to determine the attractiveness of yeast and ammonium carbonate to house flies, and whether combining and/or aging these ingredients woul d affect their attractiveness. Using this information, I would be able to make a few genera l conclusions as to where commercial fly traps and the tested attractants fit into an IPM program for house flies.

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27 CHAPTER 2 MATERIALS AND METHODS Laboratory Rearing of the House Fly House fly larvae, M. domestica Horse Teaching Unit (HTU) strain established in 2004, from Gainesville, FL, were reared in plas tic tubs (33.50 by 28.41 cm) on medium containing 250 ml of Calf Manna (Manna Pro Corp., St. Louis, MO), 1.5 liters of water, a nd 3 liters of wheat bran. The pupae were separated from the me dium by water floatation on the seventh day and dried on paper toweling. The pupae were plac ed in screened rearing cages (40.64 by 26.67 by 26.67 cm) for emergence. The screened rearing cages were screened on five sides with the sixth side used as an entryway that was covered with surgical stocking net that could be tied off to prevent house fly escape. Adult house flies were provided granulated sugar, powdered milk, and water ad libitum (Hogsette et al. 2002). Both larvae and adults were held on a 12:12 (L:D) photoperiod, 26.2 + 0.5 C, and 51.2 + 3.5% RH. Adults emerged in the cage following three days as pupae. Oviposition was induced with spent larval medium at around ten days post eclosion. The eggs collected were used to sust ain the colony. Males and females were held together in the same cages. Adult house flies were aspirate d from screen cages with a modified hand-held vacuum. Aspirated house flies were placed into a freezer (-30 C) until inactive (~1-5 min). After removal from the freezer, the house flies were pl aced on a chilled aluminum tray. House flies were counted and sexed (25 males, 25 females) and held in a deli cup (236.58 ml) for about one hour. House flies used in the design experiment and the aged attractant combination test were between three and seven days post-eclosion. Ho use flies used in scatter and attractant component experiments were between three and five days post-eclosion.

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28 Fly Trap Design Experiments Scatter Experiment An open release cage study was deve loped to evaluate the house flies’ propensity to escape from an enclosed space in a 24 hour period. Two di fferent trap designs were utilized no trap, designated the laboratory room, and a CDC (Cente r for Disease Control) net trap (Fig. 2-1). Each release cage (28.8 by 26.1 by 39.1 cm high) c onsisted of a screen cage fitted with a stocking net opening. A cotton-filled plastic cup (88.72 ml) soaked with 50 ml of 10% aqueous sugar solution was placed in releas e cage. House flies were then released from the deli cup into each release cage. A continuous channel was created by fitting a yellow funnel that was removed from a Trap n’ TossTM (Farnam Companies, Inc., Phoe nix, AZ) fly trap. The stocking net opening of the release cage wa s made continuous with the cone One half of the release cages were oriented so that th e cone opening was below the cage, while the other half were made so that the cone opening was above the release cag e (Fig. 2-2 and 2-3). Fo r the release cages that were left open to the laboratory room, af ter 24 h, the sugar water was removed, the cone removed, and the release cage was tied off for 24 h to kill the house flies remaining in the release cage. The dead house flies were counted and reco rded to determine the percentage of house flies that escaped into the laboratory room by taking the difference of how many house flies were left in the release cage as opposed to how many were in itially released. For those release cages fitted with CDC net traps, the CDC net traps were taken from the release cage, tied off and left to sit for 24 h to kill the flies. The flies were then counted, sexed, and recorded to determine the percentage of male, female, and total house flies that were tra pped in the CDC net traps. A replication consisted of a cage for top-entry or bot tom-entry treatments. Four replications were of laboratory room catch at the sa me time in the laboratory (22.8 0.02o C) on a 24:0 (L:D) photoperiod. The same procedure was done fo r the experiments with CDC net traps.

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29 Commercial Fly Trap Experiment Fly traps A comparison of six commercial fly traps was c onducted in the laboratory. Fly traps used were categorized as either bottom-entry or topentry. Bottom-entry fly traps were designed with an inverted funnel leading into a collection container. Bottom-e ntry fly traps included Trap n’ TossTM, The Advantage Fly TrapTM (Advantage Traps, Inc., Co lumbia, SC), and BC 1752 Dome (McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK ) (Figs. 2-4, 2-5, and 2-6) Top-entry fly traps were containers fitted w ith entrance(s) on the lid. Rescue! Reusable Fly Trap (Sterling International, Inc., Libe rty Lake, WA) and Victor Fly Magnet Trap (Woodstream, Lititz, PA) each had four entrance holes, while the Fly Terminator Pro (Farnam Companies, Inc., Phoenix, AZ) had only one en trance hole (Figs. 2-7, 2-8, and 2-9). The diameter for each entrance/exit hole were measured and used to calculate area. The areas were used to calculate the entrance:exit ratios for each fly trap. The slope of the funnel entrance and the volume of each fly trap’s capacity were also measured. Each entrance color was observed and recorded. Bioassay A fly attractant mixture cons isting of 5 g yeast, 0.12 g ammonium carbonate, and 75 ml of water was placed into each fly tr ap immediately after mixing. A co tton-filled plastic cup soaked with 50 ml of 10% aqueous sugar solution was placed in release cage. House flies were released from the deli cup into release cage. The stoc king net opening of the release cage was made continuous with the opening of the fly trap, which were either pl aced on top of the release cage (bottom-entry fly traps) or suspended below the release cage (top-entry fly traps) (Figs. 2-10 and 2-11). The stocking net was pulled over the set fly trap and tied off to prevent house flies from escaping between the release cage and the fly trap.

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30 After 24 h, each fly trap was removed from the release cage, placed into a sealable plastic bag, then refrigerated at 3o C for ~ 24 h. Upon removal from the refrigerator, captured house flies were extracted from the attractant mixtur e using a colander, placed on a chilled aluminum tray, counted, and sexed. The experiment was a completely randomized design comparing each fly trap’s capture efficacy. Each trial was run in the laboratory (22.8 0.02o C) on a 24:0 (L:D) photoperiod. The locations of the fly traps were randomized to minimize position e ffect. Testing included one of each fly trap (bottom-entry and top-entry) examined five times, except Fly Terminator Pro which was left out once due to availability. On two additional occasions, bottom-entry traps were tested using two Trap n’ TossTM fly traps, one Advantage Fly TrapTM, and one BC 1752 Dome Trap. Attractant Experiments Aged Attractant Experiment The Trap n’ TossTM fly trap was used as a standardized fly trap to evaluate the effect of attractant age on fly capture. Sets of three groups of fly attractants were prepared separately using 5 g yeast, 0.12 g ammonium ca rbonate, and 75 ml of water. These mixtures were held for 0, 1, 2, 3, or 4 d at 22.8 0.02o C yielding five age classes of at tractant. Following the aging of each attractant, the attractants were introduced into individual fly traps. A cotton-filled plastic cup containing 10% aqueous sugar solution and th e 50 house flies were intr oduced as previously described. The fly trap was s ecured on the cage, as before. Following the 24 h experiment, house flies were collected, counted, and sexed as in commercial trap experiments. Three cages of each attractant age mixture we re run simultaneously (22.8 0.02o C) on a 24:0 (L:D) photoperiod. The experiment was run twice. Due to contamination of release cage with insecticide, one cage of 1-d aged attractant was discarded.

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31 Attractant Component Experiment This study consisted of four treatments in cluding 75 ml water cont rol, 0.12 g ammonium carbonate, 5 g of yeast, or a combination of the ammonium carbonate and yeas t, each in 75 ml of water. Each of the attractants was aged 0, 2, or 4 d prior to placement into a Trap n’ TossTM fly trap. A cotton-filled plastic cup containing 10 % aqueous sugar solution and the 50 house flies were introduced, then the stocking net used to secure the fly trap on the cage, as before. House flies were collected, counted, and sexed as in co mmercial trap experiments. Each combination was run simultaneously and was r un independently six times, for a total of six replications. Experiment was run in the lab (22.8 0.02o C) on a 24:0 (L:D) photoperiod. Data Analysis Actual means are presented in tables. Data for percentage catch of males, females, and total for laboratory room, CDC net trap, each commercial trap, and each aged attractant were arcsine square root transformed and analyzed usi ng one-way analysis of variance. Male, female, and total percent house fly catch was analyzed by linear regression for the attractant mixture aged 1-4 days. A one-way analysis of varian ce was run on each attractant component by days aged (0, 2, 4). Means for total, male, and fema le percent catch were separated using Student Newman-Keuls (SNK) Test (P = 0.05) (SAS 2001). A Student’s t-test (P = 0.05) was used to compare the entrance position variable for labo ratory room versus CDC net trap, the catch variable for laboratory room vers us CDC net trap catch, and the sex variable for the different ages and combinations of attractants.

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32 Figure 2-1. CDC Net Trap used to capture house flies escaping from screen cages.

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33 Figure 2-2. Bottom-entry release cage design fitted with a Trap n’ TossTM cone and open to laboratory room.

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34 Figure 2-3. Top-entry release cage design fitted with modified Trap n’ TossTM and open to laboratory room

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35 Figure 2-4. Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ ) fly trap

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36 Figure 2-5. The Advantage Fly TrapTM (Advantage Traps, Inc., Columbia, SC)

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37 Figure 2-6. BC 1752 Dome (McPha il) Trap (AgrisenceTM Agriculture, Pontypridd, UK)

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38 Figure 2-7. Rescue! Reusable Fly Trap (Ster ling International, Inc., Liberty Lake, WA)

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39 Figure 2-8. Victor Fly Magnet Trap (Woodstream, Lititz, PA)

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40 Figure 2-9. Fly Terminator Pro (Far nam Companies, Inc., Phoenix, AZ)

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41 Figure 2-10. Bottom-entry study design with Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ) fly trap attached.

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42 Figure 2-11. Top-entry study design with Victor Fly Magnet Trap (Woodstream, Lititz, PA) attached.

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43 CHAPTER 3 RESULTS Fly Trap Design Experiments Scatter Experiment When the house flies were released from the de li cup into the release cage, they exhibited resting behavior preceded by a s hort time of flying or walking. They did not show any initial observational attraction to the tra p, sugar water, or opening of th e release cage. However, over half (53.38 4.18%) were eventually captu red. The cone allowed house flies to easily leave the release cage, but made it difficult to reenter. The percentage of house flies both captured in CDC net traps and escaped into the laboratory room after 24 h was determined (Table 3-1). For the released male house flies, there was no significant difference between bottom-en try and top-entry treatments (df = 9.11, t = 2.08, P = 0.0669). There was no signifi cant difference in house fly escape between bottom-entry (82.00 6.63%) and top-entry (90.00 2.58) treatm ents for the laboratory room. However, significantly more males escaped into the laborato ry room than escaped into CDC net traps (df = 9.34, t = 4.39, P = 0.0016). Significantly more ma les escaped through t op-entry (67.00 2.52%) treatments than into the bottom-entry (22.00 3 .46%) treatments into CDC net traps (df = 3, F = 22.13, P < 0.0001). Within the bottom-entry treatmen t, significantly more males escaped into the laboratory room (82.00 6.63%) than were caught in the CDC net trap (22.00 3.46%) (Table 3-1). Significantly more male house f lies escaped into the la boratory room (90.00 2.58%) than into the CDC net trap (67.00 2.52%) within th e top entry treatment. For the released female house flies, ther e was no significant difference between bottomentry and top-entry treatments (df = 14, t = 0.20, P = 0.8457). Regardless of treatment, significantly more female house flies escaped into the laboratory room than were captured in

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44 CDC net traps (df = 8.15, t = 6.43, P = 0.0002). There was no signi ficant difference in house fly escape between bottom-entry (67.00 11.12%) a nd top-entry (65.00 11.24%) treatments for the laboratory room escape option. Overall, fe w female house flies were captured in the CDC net traps (17.00 2.97%) and there was no signi ficant difference between bottom-entry (19.00 2.52%) and top-entry (15.00 3.42 %) treatments. Within the bottom-entry treatment, significantly more females es caped into the laboratory r oom (67.00 11.12%) than were captured in the CDC net trap (19.00 2.52%). Significantly more female house flies also escaped into the laboratory room (65.00 11.24%) than into th e CDC net trap (15.00 3.42%) for the top-entry treatment (df = 3, F = 11.50, P = 0.0008). There was no significant difference between th e total numbers of hous e flies that escaped through top entry and bottom entry treatments (df = 14, t = 0.90, P = 0.3844). Overall, house flies were not caught in CDC net tr aps as often as they escaped in to the laboratory room (df = 14, t = 7.67, P < 0.0001). About three quarte rs of the released house flies escaped into the laboratory room (76.00 4.04%). Top-entry CDC net traps (41.00 2.89%) caught si gnificantly more house flies than the bottom-entry CDC net traps (20.50 2.87%) (df = 3, F = 27.49, P < 0.0001). There was no significant differe nce between bottom-entry (74.50 6.99 %) and top-entry (77.50 5.06%) traps. The mean trap catch for th e CDC net traps was only one-third (30.75 4.31%) of the total released house flies. However, significantly more house flies escaped through bottom-entry traps into laboratory room (74.50 6.99%) than were captured in the bottom-entry CDC net trap (20.50 2.87%) (df = 3, F = 27.49, P < 0.0001). This pattern was also observed with top-entry traps where sign ificantly more house f lies escaped into the laboratory room (77.50 5.06%) than into the CDC net trap ( 41.00 2.89%) (df = 3, F = 27.49, P < 0.0001).

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45 Commercial Fly Trap Dimensions Six commercial fly traps were compared for their ability to captur e house flies released into screened cages (Table 3-2). Each fly trap consisted of a plastic cont ainer that held both the attractant and the captured house flie s. All fly traps had funnels that had both entrance and exit holes that facilitated house fly capture but limited escap e. Every fly trap, except the Terminator Pro, had entrance holes that were la rger than the exit holes. The Trap n’ TossTM, a bottom-entry trap, had the greatest entran ce area of all the fly traps (176.24 cm2), whereas the Fly Terminator Pro had the largest entran ce area for the top-entry fly traps (20.51 cm2). The smallest entrance area of the top-entry fly traps was the Rescue! Reusable Fly Trap (3.16 cm2), whereas the smallest bottomentry area was The AdvantageTM Fly Trap (46.57 cm2). The smallest exit area of the top-entry fly traps was the Rescue! Reusab le Fly Trap (2.00 cm2), whereas the bottom-entry fly trap with the sm allest exit area was The Advantage Fly TrapTM (9.08 cm2). The largest exit area of the top-entry fly traps was the Fly Terminator Pro (20.51 cm2), whereas the Trap n’ TossTM had the largest exit area (11.95 cm2) of bottom-entry traps. The fly trap entrance to exit ratio is a measure of the entrance area divided by the exit area. This is an important measurement because it prov ides the best relative combination of allowing house flies into the fly trap without releas ing them. The bottom-entry Trap n’ TossTM had the greatest entrance:ex it ratio of the fly traps (14.75), wher eas the Fly Magnet had the greatest entrance:exit ratio of the top-en try fly traps (2.05). The top-en try Fly Terminator Pro had the smallest entrance:exit ratio of all the fly traps (1.00), whereas BC 1752 Dome had the smallest entrance:exit ratio of the bottom-entry fly traps (4.64). The slope of the entrance cone directs the house flies into the hol ding container and the range of slopes observed by these traps was co nsiderable. The BC 1752 Dome had the greatest slope of the bottom-entry fly traps (75.57o), whereas the Terminator Pro, a top-entry fly trap,

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46 had the greatest slope of all the fly traps (90.00o). The Trap n’ TossTM, a bottom-entry fly trap, had the least slope of all the fly traps (61.67o), whereas the Rescue! had the least slope of the top-entry fly traps (85.07o). The bottom-entry fly trap with the larg est fly holding volume was Trap n’ TossTM (1936 ml), whereas the fly trap with the largest vol ume was the top-entry Fly Terminator Pro (3786 ml). The fly trap with the smallest volume was the BC 1752 Dome, whereas the top-entry fly trap with the smallest volume was the Fly Magnet (983 ml). Each fly trap’s entrance was one of several colors. The Trap n’ TossTM and BC 1752 Dome’s entrances were colored yellow. The Advantage Fly TrapTM’s entrance was white. The Fly Magnet and Fly Terminator Pro had blac k entrances, while the Rescue! had a green entrance. Commercial Fly Trap Experiment When house flies were released from deli cups into the release cag e, they showed the previously described initial behavior. At the completion of the 24 h experiment, some of the house flies were attracted to or wa ndered into the fly traps. Howe ver, it is likely that some may have escaped back into the release cage. House f lies would continue to sh ow this basic behavior throughout the experiments. The percentage of house flies captured in six different commercial fl y traps after 24 h was recorded (Table 3-3). For this experiment, the mean catch of released males was 36.67 3.17% (n = 42). The highest capture was with the The Advantage Fly TrapTM (51.43 7.33%), while the lowest catch was with the Fly Terminat or Pro (22.00 13.71%). These were the only significantly different catch means with respect to released male house flies (df = 5, F = 2.87, P = 0.0303). There was no significantly superior bottom-entry fly trap when compared only to the other bottom-entry traps or top-en try fly trap when compared only to other top-entry traps.

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47 The mean catch of released females for th is experiment was 41.90 3.56% (n = 42). The Trap n’ TossTM caught significantly more released fe males (66.22 5.89%) than the Advantage (35.43 6.32%), BC 1752 Dome (34.86 6.97%), Fly Magnet (32.80 7.31%), and Fly Terminator Pro (28.00 5.42%) (df = 5, F = 4.58, P = 0.0031). Rescue! (58.40 12.24%) was not significantly different th an the best or the worst fly traps when catching female house flies. There was no significant difference between the top-entry fly traps; however, the Trap n’ TossTM caught significantly more house flies than the other two botto m-entry fly traps (df = 5, F = 4.58, P = 0.0031). The mean capture for all fly traps in th e experiment was 39.33 2.58% of the total released house flies (n = 42). The fly trap that captured the mo st house flies was the Trap n’ TossTM (52.22 5.13%), while the Fly Terminator Pro captured the fewest (25.00 8.10%). These were the only significantly different catch means with respec t to total house flies released (df = 5, F = 3.46, P = 0.0134). There were no signifi cantly superior fly traps when comparing only top-entry fly traps with each other and bottom-entry fly traps with each other. Attractant Experiments Aged Attractant Experiment The percentage of house flies after captured after 24 h in fly traps with a yeast and ammonium carbonate mixture aged 0 4 days (by one day intervals) was determined (Table 3-4). A significant difference between aged attractants was observed with male house flies (df = 4, F = 3.44, P = 0.0233). Attractants aged for four da ys (62.00 5.54%) and three days (60.67 7.04%) caught significantly more house flies than 1-day aged attractant (35.20 5.43%). The freshly mixed attractant (38.22 3.49%) and 2-da y aged attractant (42.00 7.85%) were not significantly different from the mixtures that cap tured the most and least house flies. Regression

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48 analysis demonstrated that as attractant aged from one to four days, male house fly catch increased linearly (y = 9.907x + 25.2, R2 = 0.9073) (Fig. 3-1). As with males, a significant difference wa s also observed between female house fly capture among the attractants (df = 4, F = 3.63, P = 0.0188). The 4-day aged attractant (78.00 5.91%) caught significantly more females than attractant aged 1 day (47.20 4.80%). Significant differences were not observed among other treatm ents. Regression analysis demonstrated that as attractant aged from 1 to 4 days, female house fly catch increased linearly (y = 11.173x + 31.2, R2 = 0.8801) (Fig. 3-1). Overall, there was a significant difference between the numbers of house flies captured using variously aged attractants (df = 4, F = 4.62, P = 0.0065). The 4 day aged attractant (70.00 4.73%) captured significantly more house flies th an attractants aged two days (49.00 6.65%) and one day (41.20 4.08%). Attractant aged three days (63.00 5.99%) caught significantly more house flies than attractant aged one day ( 41.20 4.08%). Freshly mixed attractant (55.33 3.13%) was not significantly different from any of the other mixtures. Regression analysis demonstrated that as attractant aged from 1 to 4 days, total house fly catch increased linearly (y = 10.04x + 30.7, R2 = 0.9827) (Fig. 3-1). Attractant Component Experiment To determine the importance of attractant age and attractant components on house fly capture, a study was conducted examining individua l and combinations of components at 0, 2, and 4 days post-mixing (Table 3-5). The fr eshly mixed combination treatment (56.00 7.93%) captured significantly more male house flies than the ammonium carbonate (20.00 6.20%) and the control treatments (9.33 3.04%) (df = 3, F = 9.31, P = 0.0005). For attractants aged both 2 and 4 days, the combination and yeast treatments captured significantly more house flies than the

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49 ammonium carbonate and contro l treatments, respectively (df = 3, F = 10.94, P = 0.0002; df = 3, F = 8.45, P = 0.0008). The freshly mixed and 4-day-old attractant, combination and yeast treatments captured significantly more female house flies than th e ammonium carbonate and control treatments, respectively (df = 3, F = 12.97, P < 0.0001; df = 3, F = 16.98, P < 0.0001). For attractants aged 2 days, the combination treatment (69.33 8.56%) ca ught significantly more house flies than the yeast treatment (40.67 8.48%), which caught si gnificantly more house flies than ammonium carbonate (18.67 6.25%) and the control treatments (12.67 7.26%) (df = 3, F = 10.21, P = 0.0003). The freshly mixed combination treatment (61.00 5.48%) captured significantly more total house flies than the yeast treatment ( 41.67 5.55%), which capture d significantly more house flies than the ammonium carbonate (22.0 0 6.85%) and the control treatments (9.67 3.32%). With both 2-day and 4-day aged attrac tants, the combination and yeast treatments captured significantly more house flies than th e ammonium carbonate an d control treatments, respectively (df = 3, F = 11.77, P = 0.0001; df = 3, 24.27, P < 0.0001). The combination (df = 10, t = 2.91, P = 0.0155) and yeast (df = 10, t = 2.56, P = 0.0283) treatments captured significantly more female hous e flies than males when aged for 4 days. All other combinations of attractant components a nd ages did not show si gnificant differences between the sexes.

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50Table 3-1. Percent escape of male, female, and tota l house flies into CDC net traps or laboratory room. % (SE)a ____________________________________________________________________________________________________________ Trapb nc Males Females Total ____________________________________________________________________________________________________________ Bottom Entry Room 4 82.00 (6.63)a 67.00 (11.12)a 74.50 (6.99)a Net 4 22.00 (3.46)c 19.00 (2.52)b 20.50 (2.87)c Top Entry Room 4 90.00 (2.58)a 65.00 (11.24)a 77.50 (5.06)a Net 4 67.00 (2.52)b 15.00 (3.42)b 41.00 (2.89)b df = 3, F = 22.13, P < 0.0001 df = 3, F = 11.50, P = 0.0008 df = 3, F = 27.49, P < 0.0001 ____________________________________________________________________________________________________________ Data were arcsine square root transformed before analysis. Means within a column fo llowed by the same letter are not significa ntly different ( = 0.05, Student Newman-Keuls test; SAS 2001). a 25 male and 25 female, 3-5 d post-eclosi on, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Fly trap rem oved from release cage following 24 h exposure. b Room = house flies allowed to escape into laboratory room. Net = house flies captured in CDC net trap affixed to cone opening c n = replications

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51Table 3-2. Characteristics of six commercially available fly traps used to evaluate house fly capture efficacy. ____________________________________________________________________________________________________________ Trapa Entrance (cm2) Exit (cm2) Ratiob Slope (deg) Volume (ml)c Color ____________________________________________________________________________________________________________ Trap n’ TossTM 176.24 11.95 14.75 61.67 1936 yellow AdvantageTM 46.57 9.08 5.13 75.33 1840 white BC 1752 Dome 49.00 10.56 4.64 75.57 748 yellow Rescue! 3.16 2.00 1.58 85.07 1060 green Fly Magnet 6.16 3.00 2.05 89.97 983 black Terminator Pro 20.51 20.51 1.00 90.00 3786 black ____________________________________________________________________________________________________________ a Bottom-entry traps were Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ), The Advantage Fly TrapTM (Advantage Traps, Inc., Columbia, SC), and BC 1752 Do me McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK ) and were placed on top of the release cage. Top-entry traps were Rescue! Reusable Fly Trap (Sterling International, Inc., Li berty Lake, WA), Victor Fly Mag net Trap (Woodstream, Lititz, PA), and Fly Terminator Pro (Farnam Co mpanies, Inc., Phoenix, AZ), a nd were placed under the release cage. b Ratio = entrance exit c Volume = number of ml holding capacity of fly trap

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52Table 3-3. Percentage of male, female, and tota l house flies caught in six commercial traps. ____________________________________________________________________________________________________________ % Catch (SE) a ____________________________________________________________________________________________________________ Trapb n Males Females Total ____________________________________________________________________________________________________________ Trap n’ TossTM 9 38.22 (5.21)ab 66.22 (5.89)a 52.22 (5.13)a AdvantageTM 7 51.43 (7.33)a 35.43 (6.32)b 43.71 (4.40)ab BC 1752 Dome 7 22.86 (4.07)ab 34.86 (6.97)b 28.86 (5.03)ab Rescue! 5 36.00 (9.38)ab 58.40 (12.24)ab 47.20 (9.73)ab Fly Magnet 5 24.80 (5.57)ab 32.80 (7.31)b 28.80 (6.09)ab Terminator Pro 4 22.00 (13.71)b 28.00 (5.42)b 25.00 (8.10)b df = 5, F = 2.87, P = 0.0303 df = 5, F = 4.58, P = 0.0031 df = 5, F = 3.46, P = 0.0134 ____________________________________________________________________________________________________________ Data were arcsine square root transformed before analysis. Means within a column fo llowed by the same letter are not significa ntly different ( = 0.05, Student Newman-Keuls test; SAS 2001). a 25 male and 25 female, 3-7 d post-eclosi on, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Fly trap rem oved from release cage following 24 h exposure. All traps contained a combination of yeast and ammonium carbonate (41.67 : 1) in 75 ml of water. b Bottom-entry traps were Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ), The Advantage Fly TrapTM (Advantage Traps, nc., Columbia, SC), and BC 1752 Dome McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK ) and were placed on top of the release cage. Top-entry traps were Rescue! Reusab le Fly Trap (Sterling International, Inc., Liberty Lake, WA), Vi ctor Fly Magnet Tra p (Woodstream, Lititz, PA), and Fly Terminator Pro (Farnam Companies, Inc., Phoenix, AZ), and were placed under the release cage

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53Table 3-4. Percentage of male, female, and total house flies captured after 24 h in Trap n’ TossTM fly traps baited with an attractant aged up to 4 d. ____________________________________________________________________________________________________________ % Catch (SE)a ____________________________________________________________________________________________________________ Attractant age (d)b nc Males Females Total ____________________________________________________________________________________________________________ 0 6 38.22 (3.49)ab 59.33 (4.78)ab 55.33 (3.13)abc 1 5 35.20 (5.43)b 47.20 (4.80)b 41.20 (4.08)c 2 6 42.00 (7.85)ab 56.00 (7.00)ab 49.00 (6.65)bc 3 6 60.67 (7.04)a 65.33 (7.06)ab 63.00 (5.99)ab 4 6 62.00 (5.54)a 78.00 (5.91)a 70.00 (4.73)a df = 4, F = 3.44, P = 0.0233 df = 4, F = 3.63, P = 0.0188 df = 4, F = 4.62, P = 0.0065 ____________________________________________________________________________________________________________ Data were arcsine square root transformed before analysis. Means within a column fo llowed by the same letter are not significa ntly different ( = 0.05, Student Newman-Keuls test; SAS 2001). a 25 male and 25 female, 3-7 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm re lease cage. Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ) fly trap rem oved from release cage following 24 h exposure. b Combination of yeast and ammonium car bonate (41.67 : 1) in 75 ml of water c Replications

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54Table 3-5. Percentage of house flie s caught in baited Trap n’ TossTM fly traps with attract ants aged up to 4 d. ___________________________________________________________________________________________________________ Male % Catch (SE) a ____________________________________________________________________________________________________________ Days ____________________________________________________________________________________________________________ Attractantb 0 2 4 ____________________________________________________________________________________________________________ Control 9.33 (3.04)c 7.33 (2.40)b 10.67 (3.82)b Yeast 34.00 (6.26)ab 45.33 (6.98)a 37.33 (5.81)a Ammonium 20.00 (6.20)bc 16.67 (5.97)b 17.33 (4.92)b Carbonate Combination 56.00 (7.93)a 46.00 (7.14)a 44.00 (7.08)a df = 3, F = 9.31, P = 0.0005 df = 3, F = 10.94, P = 0.0002 df = 3, F = 8.45, P = 0.0008 ____________________________________________________________________________________________________________ Data were arcsine square root transformed before analysis. Means within a column fo llowed by the same letter are not significa ntly different ( = 0.05, Student Newman-Keu ls test; SAS 2001). n = 6 a 25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm re lease cage. Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ) fly trap re moved from release cage following 24 h exposure. b Control = 75 ml tap water, yeast = dried active baker’s yeas t in 75 ml of tap water, ammonium carbonate = ammonium carbonate in 75 ml of tap water, combination = yeast and amm onium carbonate (41.67 : 1) in 75 ml of tap water

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55Table 3-5 Continued. ____________________________________________________________________________________________________________ Female % Catch (SE) a ___________________________________________________________________________________________________________ Days __________________________________________________________________________________________________________ Attractantb 0 2 4 ____________________________________________________________________________________________________________ Control 10.00 (4.70)b 12.67 (7.26)c 15.33 (3.78)b Yeast 49.33 (6.67)a 40.67 (8.48)b 61.33 (7.35)a Ammonium 24.00 (8.20)b 18.67 (6.25)c 24.67 (5.79)b Carbonate Combination 66.00 (5.03)a 69.33 (8.56)a 77.33 (8.98)a df = 3, F = 12.97, P < 0.0001 df = 3, F = 10.21, P = 0.0003 df = 3, F = 16.98, P < 0.0001 ____________________________________________________________________________________________________________ Data were arcsine square root transformed before analysis. Means within a column fo llowed by the same letter are not significa ntly different ( = 0.05, Student Newman-Keuls test; SAS 2001). n = 6 a 25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm re lease cage. Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ) fly trap re moved from release cage following 24 h exposure. b Control = 75 ml tap water, yeast = dried active baker’s yeas t in 75 ml of tap water, ammonium carbonate = ammonium carbonate in 75 ml of tap water, combination = yeast and amm onium carbonate (41.67 : 1) in 75 ml of tap water

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56Table 3-5 Continued ____________________________________________________________________________________________________________ Total % Catch (SE)a ____________________________________________________________________________________________________________ Days ____________________________________________________________________________________________________________ Attractantb 0 2 4 ____________________________________________________________________________________________________________ Control 9.67 (3.32)b 10.00 (4.41)b 13.00 (3.09)b Yeast 41.67 (5.55)a 43.00 (7.30)a 49.33 (5.77)a Ammonium 22.00 (6.85)b 17.67 (5.85)b 17.67 (3.56)b Carbonate Combination 61.00 (5.48)a 57.67 (6.52)a 60.67 (5.90)a df = 3, F = 13.15, P < 0.0001 df = 3, F = 11.77, P = 0.0001 df = 3, F = 24.27, P < 0.0001 ____________________________________________________________________________________________________________ Data were arcsine square root transformed before analysis. Means within a column fo llowed by the same letter are not significa ntly different ( = 0.05, Student Newman-Keu ls test; SAS 2001). n = 6 a 25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm re lease cage. Trap n’ TossTM (Farnam Companies, Inc., Phoenix, AZ) fly trap re moved from release cage following 24 h exposure. b Control = 75 ml tap water, yeast = dried active baker’s yeas t in 75 ml of tap water, ammonium carbonate = ammonium carbonate in 75 ml of tap water, combination = yeast and amm onium carbonate (41.67 : 1) in 75 ml of tap water

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57 females y = 11.173x + 31.2 R2 = 0.8801 total y = 10.04x + 30.7 R2 = 0.9827 males y = 9.907x + 25.2 R2 = 0.90730 10 20 30 40 50 60 70 80 90 012345Attractant Age% Catch Males Females Total Figure 3-1. Effect of attractant age on percent catch of male, female, and total house flies.

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58 CHAPTER 4 DISCUSSION AND CONCLUSIONS Fly Trap Design Experiments House flies tend to scatter from sites where they are released (Hindle 1914, Jones et al. 1999). Sacca (1963) concluded that house flies disp erse in a more or less random pattern until suitable conditions are found. However, Bishop p and Laake (1919) found that house flies would pass up favorable feeding and breeding places in the c ourse of dispersal. In my experiments, the house flies released into a cage sc attered from the cage into the laboratory room and various fly traps. This scattering behavior was in accordan ce with the past literature. The house flies scattered from the release cage, which containe d nutrition, water, and po tential mates. Sacca’s (1963) observation that house flies seem to move from one place to another in a random fashion was true of the observations in my experiment s. However, Sacca (1963) stated that these random movement episodes end when suitable cond itions are found i.e. food and mates, but the behavior of the house flies in my experiments to forgo suitable conditions contradicted this. Instead the house flies in my experiments beha ved more in accordance with the findings of Bishopp and Laake (1919). This behavior may account for why more house flies escaped from the release cage into the laboratory room than the CDC net trap and the un-baited Trap n’ TossTM, because there is a small chance that th e house flies would move back towards the opening of the release cage and go back in, but as th e house flies scatter into one of the fly traps, they can just as well scatter back into the release cage. However, despite the seemingly random scatte r from the release cages, air movement may also affect the house fly dispersal behavior. Several authors have found house flies to be positively anemotropic (Hindle 1914, Hindle and Merriman 1914, Ball 1918, Carment 1922, Uvarov 1931). In my experiments, more house f lies scattered into the laboratory room than

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59 ended up in the CDC net trap, and more were in th e CDC net trap than in the un-baited Trap n’ TossTM. There is a direct correla tion with allowed air movement in this situation. The bottomentry CDC net trap versus the top-entry CDC net trap difference in house fly catch suggest the position of the trap was very important to return dispersal. For each treatment, there was the same entrance/exit cone from the Trap n’ TossTM, but the Trap n’ TossTM trap allowed less air flow than did the CDC net trap, and the CDC net trap allowed less air flow than did the open laboratory room. The behavior of the house flie s in my experiments was in accordance with previous authors’ findings, that house flies will seek and fly upwind. Most of the commercial fly traps were not signif icantly different in their abilities to capture house flies. However, the Trap n’ TossTM caught significantly more house flies than the Terminator Pro, and this was probably due to a number of design factors including large entrance hole area, small exit hole area, the ratio of entrance to exit hole ratio, the slope being close to 60o, and the color of the entrance. This is not surprising when considering the overall design of each fly trap. Picken s (1995) stated that the larger the entrance hole, the more house flies that would be captured with a fly trap. His optimum was an entrance of 506.45 cm2. I found the Trap n’ TossTM to be only ~35% of that area, but the Fly Terminator Pro was a mere 4% of Pickens’ (1995) ideal entrance area. So, it can be said that a large entrance may be partially responsible for the Trap n’ TossTM having a higher percent catch of the house flies than Terminator Pro. Pickens (1995) al so states that a small exit w ill work best to keep captured house flies in the fly trap. Hi s ideal exit area is 0.28 to 1.33 cm2. I found the Trap n’ TossTM to be almost nine times larger than 1.33 cm2, but Fly Terminator Pro is over 15 times larger. The smaller exit area of the Trap n’ TossTM may have contributed to its higher house fly capture through retention of captured house flies. If a large entrance and a small exit are best for

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60 capturing house flies, then, within reason, the high er the ratio of entrance to exit, the more house flies a fly trap should catch. Again the Trap n’ TossTM ratio was almost 15 times larger than that of the Terminator Pro. The slope may ha ve been a factor in the Trap n’ TossTM capturing more house flies than Terminator Pro. According to Pi ckens (1995) the slope of the cone of the fly trap should be 60o. The Trap n’ TossTM had a slope barely above 60o, while Fly Terminator Pro was 90o. There have been many studies documenting whic h color makes a surfac e most attractive to house flies, and there have been varied conclu sions. Awati and Swaminath (1920) used colored paper and found yellow to be most attractive. Bu rg and Axtell (1984) painted plastic jug traps and found yellow to be the most a ttractive to house flie s. Pickens (1995) stated that white or yellow were preferred in cool weather, while red, black and bl ue were the preferred color surfaces when the weather was hot, while M uniz (1967) found that color preference was independent of temperature. Hecht et al. ( 1968) using colored pieces of cardboard found that black was most attractive in the laboratory, wh ile yellow was most attr active outdoors. Nava (1967) found that house flies were at tracted to black cardboard more than any other color. The studies of Awati and Swaminath (1920), Burg and Ax tell (1984), and to some extent Hecht et al. (1968) and Pickens (1995) substa ntiate that the yellow color ma y have played a role in the yellow entrance of Trap n’ TossTM capturing more house flies than the black entrance of Fly Terminator Pro. The commercial fly traps had many different ch aracteristics, but only two fly traps were significantly different from each other. The Trap n’ TossTM entrance area was almost nine times larger and the exit area was about half as large as the Fly Termin ator Pro. The ratio then of entrance area to exit area for the Trap n’ TossTM was almost 15 times more than Terminator

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61 Pro. The slope of the entrance of Trap n’ TossTM was nearly the ideal 60o, while Fly Terminator Pro was ~30o steeper. The color of the Trap n’ TossTM entrance was yellow, whereas the Fly Terminator Pro entrance was black. Although there was no one apparent overriding characteristic that allowed the Trap n’ TossTM to capture more house flies than the Terminator Pro, these studies document the va riety of issues impacting house fly trapping. Attractant Experiments Two commonly used materials in commercial fl y trap attractants are yeast and ammonium carbonate. Pickens (1995) devise d a categorizing system for the different odor attractants for house flies. They are sugars and fermentation pr oducts, proteins, and animal excretions. Yeast is in the sugar and fermentation products ca tegory, while ammonium carbonate is in both the sugar and fermentation products as well as an imal excretions category (Pickens 1995). Cohen et al. (1991) used fly traps baited with yeast around a military camp and reported a 64% decrease in house fly density that accounted fo r a 42% decrease in clinic visits by the soldiers for diarrhea and an 85% reduction in Shigellosis visits. Ch apman et al. (1998) found that yeast provided sligh tly increased house fly attraction over th e control, while adding yeast to (Z)-9-tricosene-impregnated targets produced a si gnificant reduction in the number of male house flies attracted. In my studi es, I found yeast to be more attr active than the wa ter control at 0, 2, and 4 days old by ~30-35%. My studies do agr ee with both authors that yeast attracts more house flies than a control, but my studies put yeast’s attractiveness somewhere between the attractancy observed by either Cohen et al. (1991) or Chapman et al. (1998). Ammonium salts have been shown to be attrac tive to various filth flies most likely due to the salts’ release of am monia. Ammonium carbonate is an ammonium salt and releases both ammonium carbonate and carbon dioxide (Loeser et al. 2004). Mulla et al. (1977) lists ammonium chloride and ammonium sulfate, as highly attractive to hous e flies. Wieting and

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62 Hoskins (1939) found ammonia attractive and ca rbon dioxide not attrac tive to house flies. Hobson (1935, 1936) found ammonium carbonate in c onjunction with indole was attractive to Lucilia sericata (Meigen). Mulla and Ridsdill-Smith (1986) found that when ammonium carbonate was added to an attractant mixture, attractancy was not sign ificantly increased for Musca vetustissima My studies demonstrated that ammonium carbonate was never more attractive than the control, and when added to y east (an attractant) did not increase attractancy. The results that I obtained were never directly comparable with the aforementioned studies due to the difference in attractant components used and/or species tested against. However, my studies agree most with Mulla and Rids dill-Smith’s (1986) experiments with M. vetustissima, in that ammonium carbonate does not represent an at tractant chemical for the house fly when used alone or in conjunction w ith other attractants. Initially, there was reason to believe that a combination of yeast and ammonium carbonate would be more attractive than the yeast or ammonium carbonate alone. Cosse and Baker (1996) used Eletctoantennagram (EAG) to determine a number of chemicals th at illicit a response in house flies, but many required combination with other attractants to stimulate a behavioral response i.e. attraction. Geden (2005) combin ed two commercial attractant mixtures and reported synergistic effects. When yeast and ammonia carbonate were mixed, the combination did not catch significantly more house flies than the yeast alone, but did catch more than ammonium carbonate alone. There was certainly no synergistic effect. Because the assumed attractive ammonium carbonate turned out to be not attractive, my results do not agree or disagree with Cosse and Baker (1996) or Geden (2005). Given the nature of the presumed attract ant combination used (yeast and ammonium carbonate components), it is logical to assume th at the attractant combination would change over

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63 time. Ammonium carbonate quickly released amm onia and carbon dioxide when added to water. The yeast was a living organism and its life cycle was subject to availabil ity of nutrition. In my experiment, there was not an ample carbon supply for the yeast i.e. sugar, meaning the yeast either arrested in G0 phase, progressed slowly through the life cycle, or died (Dickinson and Schwiezer 1999). How the yeast changed is unknow n, but it was changing. Pickens et al. (1973) stated that baits of fermented grains and yeas ts took about three to four days to become attractive. Geden (2005) observe d commercial attractants to beco me less attractive by the fourth day. The attractant combination in my study increas ed linearly from day 1 to 4 and agrees more with Pickens et al. (1973), most likely due to the similarity of fermentation product attractants that we each used. The components of the at tractant mixtures used by Geden (2005) were not reported; therefore it is difficult to discuss the similarities and differences. Also the experiments of Geden (2005) were run outdoors at temperatures as high as 37o C, which may have affected volatility or stability of the attractants, while my attractants were aged in less extreme temperatures indoors (22.8 0.02o C) which may have kept the yeast viable. Attractants increase the effectiveness of fly traps consid erably. In fact, Kuzina (1940) stated that odor was the most important mechan ism used by house flies to locate food sources. In my studies, yeast proved to be much better at attracting house flies than ammonium carbonate, and ammonium carbonate never attr acted significant numbers of house flies. The fact that ammonium carbonate was not attract ive, resulted in the combin ation of yeast and ammonium carbonate being no better at attrac ting flies than the yeast alone. When using a live organism such as yeast in order to attract house flies, as long as it is sustained, the attraction will increase linearly for at least 4 days.

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64 Conclusions House flies are important pests that are best controlled by using IPM practices. This is particularly true given the house fly’s ability to develop insecticide resistance and its considerable population potential (Hodge 1911, Sc ott et al. 2000, Learmount et al. 2002, Gao and Scott 2006). Commercial fly traps have two potential niches in a house fly IPM program, surveillance and control. Field trials must still be run in order to obtain a better understanding on how well the tested commercial fly traps perform in non-laboratory conditions. However my laboratory studies demonstrated that fly traps can be viable de vices for catching house flies. Although the Fly Terminator Pro with an un-ag ed attractant mixture of yeas t and ammonium carbonate only captured ~25% of the total hous e flies, the Trap n’ TossTM with the same attractant mixture aged 4 days, caught ~70% of the total house flies, a nd almost 80% of the female house flies. A capture of ~70% rivals the effectiveness of some biological control agen ts and insecticides. Aside from comparing with other control meas ures, ~70% capture exceeds Dadour and Cook’s (1992) requirement that in order to reduce a house fly population by 50-90%, 24-58% of the present house flies should be captured. According to my labor atory studies, the proper design (e.g. at least a 176.24cm2 entrance area, at most an 11.95 cm2 exit area, a ratio of at least 14.75, a slope around 60o, and a yellow entrance) and an aged at tractant can capture high numbers of house flies and should be considered for use as a control measure in an IPM program. In addition to controlling fly populations, commercial fly traps may serve as monitoring tools in a house fly IPM program. Burg and Axte ll (1984) suggested that fly traps represented a simple, practical way to monitor house fly populations, t hus giving a better idea of when to use additional control measures. Therefore, the comm ercial fly traps that did not catch the most house flies in these studies could still be important tools in house fly IPM.

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65 In conclusion, my studies have shown that commercial fly traps have potential as house fly IPM tools. They are versatile devices to be us ed in IPM programs, and can be used either as control devices or monitoring house fly populations. They are compatible with both insecticides and biological control measures. Also, if there exists a situa tion in which insecticides or biological controls cannot be use d, they are safe, easy, and effective enough to be used as standalone devices. According to my laboratory studies, and with th e further exploration with field studies, commercial fly traps repr esent a viable tool for managing house flies in a wide variety of settings including agriculture, urban, and military

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66 LIST OF REFERENCES Adams, T. S., and G. G. Holt. 1987. Effect of pheromone components when applied to different models on male sexual behavior in the housefly Musca domestica Journal of Insect Physiology 33:9-18. Appling, J. 1988. Toxicology on tap: Uncertain health significan ce of contaminated; drinking water. New York's Food & Life Sciences Quarterly 18:24-26. Awati, P. R., and C. S. Swaminath. 1920. Bionomic s of houseflies. III. A preliminary note on attraction of houseflies to certa in fermenting and putrefying su bstances. Indian Journal of Medical Research 7:560-567. Axtell, R. 1970a. Fly control in caged-poultry ho uses: Comparison of larviciding and integrated control programs. Journal of Economic Entomology 63:1734-1737. Axtell, R. 1970b. Integrated fly-control progra m for caged-poultry houses. Journal of Economic Entomology 63:400-405. Ball, S. 1918. Migrations of insects to Reb ecca Shoal Light-Station and the Tortugas Islands, with special reference to mosquitoes and f lies. Papers of the Department of Marine Biology Carnegie Instit ute Washington 12:193-212. Basden, E. B. 1947. Breeding the house-fly ( Musca domestica L.) in the laboratory. I. Introduction and technique. Bulletin of Entomological Research 37:381-387. Bishopp, F., and L. Henderson. 1946. Hous efly control. Usda Leaflet 182. Bishopp, F., and E. Laake. 1919. The dispersion of flies by flight. Journal of Economic Entomology. 12: 210-211. Bishopp, F., and E. Laake. 1921. Dispersion of flies by flight. Journal of Ag ricultural Research 21:729-766. Bishopp, F. C., W. E. Dove, and D. C. Parm an. 1915. Notes on certain points of economic importance in the biology of the house fl y. Journal of Economic Entomology 8:54-71. Blair, D. M. 1945. Baited fly traps. R hodesia Agricultural Journal 42:488-492. Bloomquist, G. J., T. S. Adams, and J. W. Dillwith. 1984. Induction of female sex pheromone production in male houseflies by ovary implants or 20-hydroxyecdysone. Journal of Insect Physiology 30:295-302. Burg, J., and R. Axtell. 1984. Monitoring house fly, Musca domestica (Diptera: Muscidae), populations in caged-layer poul try houses using a baited jug-trap. Environmental Entomology 13:1083-1090.

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76 Wieting, J., and W. Hoskins. 1939. The olfactory response of flies in a new type of insect olfactometer. II. Responses of the hous efly to ammonia, carbon dioxide, and ethyl alcohol. Journal of Economic Entomology 32:24-29. Wills, L., B. Mullens, and J. Mandeville. 1990. E ffects of pesticides on filth fly predators (Coleoptera: Histeridae, Sta phylinidae; Acarina: Macrocheli cidae, Uropididae) in caged layer poultry manure. Journal of Economic Entomology 83:451-457. Zimgrone, L. D., W. N. Bruce, and G. C. Decker. 1959. A mating study of female house fly. Journal of Economic Entomology 52:236. Zurek, L., S. Denning, C. Schal, and D. Watson. 2001. Vector competence of Musca domestica (Diptera: Muscidae) for Yersinia pseudotuberculosis Journal of Medical Entomology 38:333-335. Zurek, L., D.W. Watson, S. B. Krasnoff, and C. Schal. 2002. Effect of the entomopathogenic fungus, Entomophthora muscae (Zygomycetes: Entomophthoraceae), on sex pheromone and other cuticular hydro carbons of the house fly, Musca domestica Journal of Invertebrate Pathology 80:171-176.

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77 BIOGRAPHICAL SKETCH Ryan Merrill Welch was born on June 23, 1982, to Dwight and Amy Welch, in Jacksonville, FL. There he attended Parkwood Heights Elementary School, Arlington Middle School, and Stanton College Preparatory School He graduated from Stanton College Preparatory School with high sc hool and international baccalau reate diplomas in 2000. He attended the University of Florida from 2000 to 2004 and graduated cum laude earning a Bachelor of Science in entomology and a Bachelor of Arts in political science. He attended the University of Florida from 2004 to 2006 and receiv ed his Master of Science in entomology under Dr. Phil Koehler’s guidance.


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Title: Relative Effectiveness of Commmercial Fly Traps and Ammonium Carbonate/Yeast in Catching House Flies (Diptera: Muscidae)
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Copyright Date: 2008

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RELATIVE EFFECTIVENESS OF COMMERCIAL FLY TRAPS AND AMMONIUM
CARBONATE/YEAST IN CATCHING HOUSE FLIES (DIPTERA: MUSCIDAE)




















By

RYAN MERRILL WELCH


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2006

































Copyright 2006

by

Ryan Merrill Welch









ACKNOWLEDGMENTS

I express deep gratitude to Dr. Phil Koehler for his direction in this thesis and in my

graduate study as a whole. His guidance and concern have been extraordinary. I also

acknowledge my graduate committee-Dr. Chris Geden, Dr. Phil Kaufmann, and Dr. Richard

Patterson-for their help in making this thesis possible with revisions and general support. Also,

thanks go to Dr. Phil Kaufmann for allowing space in his laboratory to rear house flies for my

experiments. Ricky Vazquez, Jeff Hertz, and Matt Aubuchon offered valuable help in rearing

house flies. Special thanks go to my friends and co-workers in the urban entomology lab,

especially Gil Marshall and Osborne "Tiny" Willis, for their help in all things related to the

urban laboratory. Debbie Hall was an invaluable help with so many aspects of my graduate

experience from registering for classes to general advice. Finally, I would like to thank my

family for their lifelong support of my academic endeavors.









TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ...............................................................................................................3

LIST O F TA B LE S ......... ..... .............. ................................................................. 6

LIST OF FIGU RE S ................................................................. 7

ABSTRAC T ...........................................................................................

CHAPTER

1 REVIEW OF LITERATURE ....................................................... 10

Distribution and Classification ................................. .......................... ......... 10
Morphology/Identification ............... .......... ...... .......... 10
L ife C y cle ..........................................................................10
Flight and D ispersal ................................... ...... ............................ 12
M eating B behavior ........................................................... .. ..................13
O v ip o sitio n ................................... ..... ........................................................... .......... 15
F e e d in g ............................................................................1 6
L o n g ev ity ......... ........... .......................... ...........................17
Sex Ratio........................ .................... .................. 18
Economic Importance ........................................................... ....... .........18
C o n tro l ............................................................................2 0
F ly T ra p s ........................................................................2 4
O bj ectives .............. ...................................................................................................26

2 M A TER IA L S A N D M ETH O D S ..................................................................................... 27

Laboratory Rearing of the House Fly ................................ ........................ ............ 27
Fly Trap Design Experiments ................................ ........................... .. ......... 28
Scatter E xperim ent ................................................................28
Commercial Fly Trap Experiment..................... ...............29
F ly trap s ........................................................2 9
B io a ssa y .............................................................................2 9
A ttractant E x p erim ents ..................................................................................................... 3 0
A ged A ttractant Experim ent ..................................30............................
Attractant Component Experim ent .................................................... 31
D ata A n aly sis ........... .... ................ ............................................................................3 1

3 R E SU L T S .............. ... ................................................................43

Fly Trap D design Experim ents ........................................43...........................
Scatter E xperim ent ................................................................43
Commercial Fly Trap Dimensions .................................................45


4









Com m ercial Fly Trap Experim ent........................................................ ............... 46
A ttractant E x p erim ents ............................................................................... ......................4 7
Aged Attractant Experiment ............................................................ ................47
A ttractant Com ponent Experim ent............................................................. ............... 48

4 DISCUSSION AND CONCLUSIONS ............................................................................58

Fly Trap Design Experiments ........................................................... .. ............... 58
Attractant Experiments .............. ..... ............................. ........... 61
C onclu sions.......... ..........................................................64

L IST O F R E F E R E N C E S ...................................................................................... ...................66

B IO G R A PH IC A L SK E T C H .........................................................................................................77









LIST OF TABLES


Table page

3-1 Percent escape of male, female, and total house flies into CDC net traps or laboratory
room ..............................................................................................50

3-2 Characteristics of six commercially available fly traps used to evaluate house fly
capture efficacy .............................. .. ................. 51

3-3 Percentage of male, female, and total house flies caught in six commercial traps............52

3-4 Percentage of male, female, and total house flies captured after 24 h in Trap n'
TossT fly traps baited with an attractant aged up to 4 d....................................... 53

3-5 Percentage of house flies caught in baited Trap n' TossTM fly traps with attractants
aged up to 4 d ............................................................................... 54









LIST OF FIGURES

Figure page

2-1 CDC Net Trap used to capture house flies escaping from screen cages............................32

2-2 Bottom-entry release cage design fitted with a Trap n' TossT cone and open to
lab o rato ry ro o m ....................................................... ................ 3 3

2-3 Top-entry release cage design fitted with modified Trap n' TossT and open to
laboratory room .............................................................................................. 34

2-4 T rap n' TossTM fly trap ...................................................................................35

2-5 The A advantage Fly TrapTM .........................................................................36

2-6 B C 1752 D om e (M cPhail) Trap............................................... .............................. 37

2-7 R escue! R usable Fly Trap ................................................. ................................ 38

2-8 V ictor F ly M agnet T rap ......................................................................... ...................39

2-9 Fly Terminator Pro.................................... ..... ............ ........... 40

2-10 Bottom-entry study design with Trap n' TossTM ................................................... 41

2-11 Top-entry study design with Victor Fly Magnet Trap.............................................42

3-1 Effect of attractant age on percent catch of male, female, and total house flies ..............57









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

RELATIVE EFFECTIVENESS OF COMMERCIAL FLY TRAPS AND AMMONIUM
CARBONATE/YEAST IN CATCHING HOUSE FLIES (DIPTERA: MUSCIDAE)

By

Ryan Merrill Welch

December 2006

Chair: P.G. Koehler
Major Department: Entomology and Nematology

House flies are important pests that are best controlled by using integrated pest

management (IPM) because of the house fly's ability to develop insecticide resistance and its

considerable reproduction potential. Commonly used IPM tactics include sanitation,

insecticides, biological control agents, and fly traps.

My studies deal with the ability of commercial fly traps to catch house flies and the

evaluation of house fly behavior in relation to fly trap catches. In a series of experiments, house

flies were released into screened cages fitted with commercial fly trap funnels. The released

house flies escaped the cage either into the laboratory room or into fly traps. However, despite

the seemingly random escape from the release cages, air movement may also affect house fly

dispersal behavior. In my experiments, more house flies scattered into the laboratory room than

were captured in the un-baited CDC net trap, and more house flies were captured in the un-baited

CDC net trap than in the un-baited Trap n' TossTM. The behavior of the house flies in my

experiments showed the house flies' tendency to fly upwind. Six commercial fly traps were

compared for their ability to capture house flies released into screened cages. The Trap n'

TossTM fly trap caught significantly more house flies than the Fly Terminator Pro fly trap, and

this was probably due to a number of design factors including a large entrance hole area, a small









exit hole area, a large ratio of entrance to exit hole, the cone slope being close to 600, and the

color of the entrance cone.

Two commonly used materials in commercial fly trap attractants, yeast and ammonium

carbonate, were tested to identify their attractancy to adult house flies released into screen cages.

In my studies, yeast was more attractive than water at 0, 2, and 4 days old by -30-35%. My

studies also found ammonium carbonate was never more attractive than water, and when added

to yeast did not increase attractancy. Therefore, ammonium carbonate was not a chemical

attractant for the adult house fly when used alone or in conjunction with the other attractants

tested. Yeast, being a living organism, had a life cycle that was subject to availability of

nutrition. The attractant combination (yeast and ammonium carbonate in a 41.67:1 ratio) in my

study increased linearly from day 1 to 4. My results suggest that when using yeast to attract

house flies, as long as the yeast is protected from temperature extremes, house fly attraction will

increase linearly for at least 4 days.

My studies have shown that commercial fly traps have great potential as house fly IPM

tools. They are versatile devices to be used in IPM programs, and can be used both as control

devices and for monitoring house fly populations. According to my laboratory studies, and with

the further exploration with field studies, commercial fly traps represent a viable tool for

managing house flies in a wide variety of settings, including agriculture, urban, and military.









CHAPTER 1
REVIEW OF LITERATURE

Distribution and Classification

The house fly, Musca domestic L., belongs to the kingdom Animalia, phylum

Arthropoda, class Insecta, order Diptera, suborder Cyclorrhapha, family Muscidae, subfamily

Muscinae (West 1951). Linnaeus described the species in 1758. Other common names include

typhoid fly, cholera fly, dysentery fly, and enteric fly after important diseases thought to be

vectored by the fly at certain times and places (West 1951).

The house fly is for all practical purposes considered worldwide in its distribution,

although it may not be found in arctic regions or places of high altitude (West 1951). The fact

remains that it is a nearly ubiquitous, synanthropic insect that is found from sub-polar regions to

the tropics (Graham-Smith 1914, West 1951).

Morphology/Identification

The house fly is a gray insect that is usually 4 to 9 mm long depending on available

resources. The wings are held directed posteriorly when at rest. There is a sharp bend in the

fourth longitudinal wing vein. There are four darkened, parallel stripes running vertically down

the thorax (West 1951). The abdomen is a gray to yellow color with a dark midline and dark

markings set irregularly. Females are usually slightly larger than males and are easily

distinguished by noting the space between the eyes, in which the females' is almost twice as

large (West 1951).

Life Cycle

The house fly, like all Diptera, is holometabolous, with distinct egg, larva, pupa, and adult

stages (Chapman 1998). The first stage in any insect development, including M. domestic, is

the egg. The eggs are white and appear polished, due to a clear, viscous coating (Newstead









1908, West 1951). They are about one millimeter in length and have a pattern of small

hexagonal markings due to the shape of the egg tube in which the ovum developed (West 1951).

The egg may hatch in as little as 7 hours and 35 minutes when the temperature is an optimum 34

to 38C and in as long as three days if the temperature is as low as 10C (Hewitt 1914, Larsen

and Thomsen 1940). The usual time to hatch is between eight and 20 hours. Eggs that become

too dry will not hatch.

Larvae, or maggots, emerge from the egg by way of a slit that appears on the dorsal side of

the egg (West 1951). The larvae are white to cream-colored and possess no eyes, legs, antennae,

or other appendages (West 1951). The larva must pass through three instars reaching 7-12 mm

before pupating (Newstead 1908, West 1951). The third instar migrates from the food source to

a dry, cool place to pupate. The larval stage can take anywhere from five days to eight weeks,

depending on the temperature and food quality (Newstead 1908).

The fly reaches the pupal stage when the movements of the contracted larvae have ceased

and it is still a cream color, and ends when the adult fly emerges (Larsen and Thomsen 1940).

The puparium color quickly changes from the cream color darkening to a reddish brown or black

(Newstead 1908, West 1951). It is usually around 6 to 7 mm long and weighs about 21 mg

(Newstead 1908, West 1951). The pupal stage typically accounts for more than half of the

immature development time of the fly. It may last anywhere from 3.75 to 28 days, depending on

the food of the larval stage and the temperature (Newstead 1908, Larsen and Thomsen 1940).

Development time is inversely proportionate to temperature to a certain threshold. The upper

temperature range in which retardation becomes apparent is between 36 and 40C (Larsen and

Thomsen 1940). If the medium reaches temperature below 10C, there is a considerable level of

mortality (Sacca 1963).









The adult, or imagio, emerges from the puparium by breaking away the anterior end with

the ptilinum found on the frontal region of the head (Newstead 1908, West 1951). The fly

escapes the puparium and escapes the pupal medium such as manure, debris, sand, etc. by

contracting the ptilinum (West 1951). This important structure is expanded and contracted by

changes in blood pressure and retraction of muscles (Laing 1935, West 1951).

The newly emerged adult crawls about rapidly as the wings unfold and the exoskeleton

becomes hard. The wings become flat, thin, and transparent by withdrawal of fluids and are

supported by the veins. At last, the ptilinum is withdrawn and only the frontal "lunule" that

marks its former location is left behind (Newstead 1908, West 1951). Overall, the whole process

from egg to adult, like every life stage, varies with temperature. However, when the medium is

33.22C, the process takes about 6.5 days (Hogsette 1995).

Flight and Dispersal

House flies are able to fly considerable distances. The extent of this distance is different

according to different authors. Bishopp and Laake (1921) found that within 24 hours of release,

house flies average a distance of about 9.66 km, with a maximum distance of about 21.15 km.

Schoof and Siverly (1954) conducted a study that suggests that flies can travel as far as 32.19 km

by way of flight. They also found that large numbers of flies might migrate up to 6.44 km.

However, most flies are not recovered at distances greater than 3.22 km. This is because house

flies have been shown to fly in a more or less random pattern, until a suitable place is located

(Sacca 1963). The house fly tends to travel shorter distances in urban areas where structures,

sources of food, and sources of oviposition are more readily available, whereas in rural

environments, the fly will have to disperse further to find suitable sites (Hindle 1914, Murvosh

and Thaggard 1966). Additionally, air temperature limits the flight ability of the house fly. As

temperature increases, so does the motility of the house fly (Sacca 1963).









Arguably the most important means of dispersal for the house fly, and almost certainly the

reason for its worldwide distribution, is hitchhiking. Flies can travel great distances by using

large mammals or human-assisted transportation (Sacca 1963).

Mating Behavior

The mating behavior of the house fly is a complicated process that, once sexual maturity is

reached, is always initiated by the male. Many researchers have investigated this issue.

However, the temperatures used in their studies were not always the same. Males require at least

16 hours to mature, whereas females require at least 24 hours (Murvosh et al.. 1964a). Chang

(1965) determined a male maturation time of at least 20 hours and female maturation time of at

least 40 hours. Despite the fact that female flies may not be ready to copulate, males persist.

This may lead to a female fending off and possible damaging of the male, especially his wings,

with her posterior legs that are held near the tip of the abdomen (Bishopp et al. 1915).

Copulation may occur much later even though males aggressively try. Bishopp et al.

(1915) found flies mating as late as 16 days after emergence. This large range of time may be, at

least in part, the result of nutrition. The nutrition may affect either the sexual development or the

desire to mate. Bishopp et al. (1915) observed mating within four days of emergence on flies fed

with milk and horse manure, while a group fed partially ripened peaches and horse manure

required 16 days. Copulation was not observed among the unfed flies.

Sexually determined males seek females using visual cues. The male is sexually

aggressive, and will attempt to mate with anything resembling a female house fly, including male

house flies, fly cadavers, and inanimate objects (Rogoff et al. 1963). However, visual cues are

not necessary for a male to locate a mate. In fact, mating can occur in complete darkness

(Basden 1947, Rogoff 1965).









The male fly also uses chemical cues to identify a partner (Patterson 1957). Both male and

female house flies have gender-specific hydrocarbons, as well as hydrocarbons common to both

sexes (Nelson et al. 1981). The major pheromone of the house fly is found on the female. It is a

C23 hydrocarbon called (Z)-9-tricosene (Carlson et al. 1971). The activity of this pheromone is

enhanced by other hydrocarbons including methylalkanes, (Z)-9,10-epoxytricosane, and (Z)-14-

tricosen-10-one (Uebel et al. 1976, Nelson et al. 1981, Bloomquist et al. 1984). It works as a

short-range attractant, sex-recognition factor, arrestant, and copulatory stimulant of males

(Mayer and James 1971, Mansingh et al. 1972, Carlson et al. 1974, Adams and Holt 1987,

Hanley et al. 2004).

When the male does locate an appropriate partner, both flies begin seemingly random

movements. The male will stop movement when he gets within a few centimeters of the female,

wherein he leaps on her back (Rogoff et al. 1963). The mount may also be attempted in the air,

but the couple quickly comes to a rest, as copulation never happens in flight. If disturbed,

however, the couple may fly to another surface together (Murvosh et al. 1964a). Mating occurs

in the superimposed position, with the male above the female and both facing the same direction.

If the male is too small or disturbed, it may face at a near 900 angle or even face the opposite

direction (Hardy 1944, West 1951). After mounting, the male caresses the head of the female

with the tarsi of his front legs. This probably has a reflex effect on the female as it usually

results in ovipositor extension until it contacts the male genital atrium. The aedeagus extends as

does the accessory copulatory vesicles of the female so that they contact each other. This is

when the spermatozoa make their way into the spermethecae of the female (West 1951). The

spermatozoa are transferred to the female in as little as three to five minutes however, the couple

will usually stay in coitus for a much longer time (Murvosh et al. 1964a). Coitus has been









observed to last anywhere from 30 minutes to two hours (Hampton 1952, Murvosh et al. 1964a,

Chang 1965).

After mating, the female usually remains monogamous, and only a few mate more than a

few times (Riemann et al. 1967). In a study by Zimgrone et al. (1959), only two percent of the

females mated multiple times. One fertilization episode is sufficient for the entire egg-laying

period (Zimgrone et al. 1959). The male seminal fluid, and not mechanical stimulation or sperm,

has been shown to cause the loss of female sexual receptivity (Riemann et al. 1967).

Oviposition

Many factors are involved in ovigenesis and oviposition. For instance, diet is of utmost

importance. Protein is necessary for the production of mature eggs in the female house fly, as

with many other dipterans (Sacca 1964). Whereas the diet is integral in the production of eggs, it

is also important insofar as how quickly or slowly the eggs are oviposited. Bishopp et al.. (1915)

provided one group of flies peaches for food as opposed to another lot being supplied with milk,

peaches, and horse manure. Those fed peaches required 16 days to oviposit, whilst those fed the

mixture of foods took five. This suggests that variety, or probably more specifically, the

presence of protein, in the diet allows the female to oviposit more quickly.

Temperature also plays an important role in the rapidity of egg deposition. In general,

within reasonable ranges, speed of egg deposition is positively correlated with temperature

(Bishopp et al.. 1915). Oviposition will altogether cease if the atmospheric temperature drops to

10 C or lower, regardless of the substratum temperature (Kalandadze and Chilingarova 1942,

West 1951).

Humidity is also a factor in time to first oviposition. As observed with temperature, when

humidity increases, the speed of egg deposition increases. The humidity may make the media is

more moist, and therefore more appealing to the female flies (Bishopp et al.. 1915). However,









because liquid medium is hardly ever a suitable location, there appears to be an upper limit to

this effect (Kalandadze and Chilingarova 1924).

Taking into account the variety of factors impacting egg deposition, a female house fly

may require from five to twenty days post-eclosion to deposit her first batch of eggs (Griffith

1908, Bishopp et al.. 1915). Once she is ready to oviposit, she must find an appropriate medium.

An appropriate medium is one that will provide ample nutrition to the larvae that will hatch from

the eggs. The larvae develop most quickly in any site that is fermenting and vegetal in nature.

Flies often oviposit in garbage and manure (Mellor, J.E.M. 1919, Kalandadze and Chilingarova

1942, West 1951, Sacca 1963, Cosse and Baker 1996). Another factor influencing oviposition is

the presence of other flies. Jiang et al. (2002) documented increased oviposition at sites where

other female flies were present, possibly due to the semiochemicals n-tricosane and (Z)-9-

tricosene. When a suitable spot is located, a female fly will orient to cracks and crevices in

where she either inserts her ovipositor, or crawls into the crevice so that the eggs are protected

from desiccation and actinic light (Newstead 1908, West 1951).

House fly reproductive capacity is considerable. Hodge (1911) calculated that if one pair

of flies, unchecked, began reproducing in April, by August there would be

191,010,000,000,000,000,000 flies. It is estimated that this would cover the surface of the earth

to a depth of 47 feet. One female fly usually oviposits anywhere from 75 to 159 eggs in batches

(Newstead 1908, Dunn 1923, Herms and James 1961). House flies have been reported to

produce between two and twenty-one egg batches at intervals of eight to fourteen days, though

usually only two to four batches are deposited (Griffith 1908, Bishopp et al. 1915, Dunn 1923).

Feeding

House flies, like all animals, require nutrition in order to maintain body processes, such as

flight, movement, feeding, etc. and to provide for the building blocks of the organs, especially









the reproductive system. For the house fly, sugars or assimilable starch is required, while

proteins or byproducts of protein hydrolysis are necessary for organ formation (Kobayashi 1934,

West 1951).

Certain substances and chemicals are feeding attractants. Although house flies are

omnivorous, there are suitable, but unattractive, foods that require stimulus for consumption.

The sugars that are particularly attractive to adult flies for feeding purposes include lactose and

dextrin. Sugars that are not necessarily attractive despite their nutritive potential are glucose,

maltose, starch, or sucrose (Richardson 1917). Richardson (1917) also found that succinic and

lactic acid were feeding stimulants. Awati and Swaminath (1920) found ammonia, hydrogen

sulfide, and phosphorous compounds to be adequate to induce feeding in the house fly adult.

Robbins et al. (1965) found casein, more specifically, its amino acid components leucinee, lysine,

isoleucine, and methionine) and the guanonsine monophosphate of yeast hydroloyzate to be

feeding stimulants.

Regardless of how attractive or nutritive a food substance is to the house fly, it can only be

consumed if it is in a liquid or semi-liquid form, or is capable of being rendered so. Some solid

foods are, if soluble, put into solution by the house fly's use of vomit from the crop. There are

some instances where non-soluble solids will be taken up, then broken down further with

prestomal teeth. After the food is ingested, it will make its way through the alimentary tract

(Sacca 1963).

Longevity

The longevity of house flies, no doubt, is directly related to the temperature and humidity.

West (1951) states that in the summer flies live no longer than a few weeks, whilst in the colder

months, three months is realistic. Many studies have examined house fly longevity. Herms

(1928) recorded a mean of 30 days. Murvosh et al. (1964b), found that at a mean of 23.9 C,









males live an average of 33 days, and females 43 days, with a range of one to 99 days.

Rockstein (1957) found that when the temperature was 26.7C and relative humidity was 45%,

male house flies lived from 15 to 40 days, and female house flies lived from 20 to 60 days.

The age of the parents affects the longevity of the offspring. The average life span of the

succeeding generations decreases if only eggs from young flies are used. Flies developing from

females aged 20 to 30 days at oviposition resulted in an increase in the average lifespan

(Callahan 1962).

Sex Ratio

Studies with lab colonies suggest that the sex ratio of house flies is about 1:1. Murvosh et

al. (1964a) reported a sex ratio of 50.6 to 49.4 ratio based on 8700 individuals, and a ratio of 53.5

to 46.5, based on 5233 individuals. Rogoff (1965) determined his sex ratio using 16 different

samples of 100 pupae each, and reported similar results.

West (1951) observed that when the average adult size is smaller than 6-7 mm in length,

then males would predominate, while when the average adult size is larger, the males never

predominate the population. Herms (1928) showed, while working with Lucilia sericata

(Meigen), that larval females require more nourishment than their male counterparts to survive to

the pupal stage. This being the case, an abundance of males may signify a lack of larval nutrition

in the area.

Economic Importance

House flies gather at garbage, compost, sewage, manure, livestock facilities, and other

unsanitary sites, and then readily enter buildings making them a considerable nuisance to people

and animals (Chapman et al. 1998). Although the house fly is not as much of a problem in many

advanced industrial societies due to personal transportation being automobiles instead of horses,

it is still one of, if not the most, important pests in markets, stores, and animal sites due mostly to









annoyance (Cosse and Baker 1996, Robinson 1996, Chapman et al. 1998, Hogsette 2003). They

become most annoying when they land on people and animals causing irritation. When they

crawl on food the potential for pathogen transmission is raised. Whilst resting on walls and

windows they leave behind specks from vomit and feces. Mere annoyance has turned into

litigation. When flies from livestock facilities find their way into the spreading suburban and

urban areas, lawsuits follow (Campbell 1993). The annoyance that the flies cause livestock may

also reduce production yields (Cosse and Baker 1996, Chapman et al. 1998).

House flies are economically important as disease vectors. House flies have been long

suspected of disease transmission (Howard 1911). However, it was not until the studies carried

out by Watt and Lindsay (1948) and Lindsay et al. (1953), that there was convincing evidence

for the house fly's involvement. After a fly has become infected with a pathogen, it can transmit

the disease one of four ways. These include contact with the body setae, setae of the feet,

regurgitation, and passage through the digestive tract (West 1951). While West (1951) and

Greenberg (1973) indicate that of these four, the passage through the digestive tract is probably

the most important because it allows for multiplication of the disease organism within the

alimentary canal, Zurek (2001) shows that the potential for a pathogen to be spread via vomitus

or feces is dependent on the specific pathogen.

Greenberg (1971) lists over 30 viruses, hundreds of bacteria species, as well as a number

of fungi, protozoa, and nematodes that are associated with M. domestic. A far from exhaustive

list of some of the disease organisms that house flies are known to transmit are Escherichia coli

(including 0157:H7), .\/ngel//l spp., Salmonella spp., Helicobacterpylori, Yersinia

pseudotuberculosis, and the organisms responsible for typhoid fever, cholera, summer diarrhea,









dysentery, tuberculosis, anthrax, opthalmia, and parasitic worms (West 1951, Greenberg 1971,

Grubel et al. 1997, Iwasa et al. 1999, Zurek et al. 2000, Geden 2005).

Control

The management of the house fly is an exercise in problem solving. West (1951) broke the

control of house flies into two categories: planned control and emergency control. This approach

is better known as integrated pest management (IPM) today. West's apparent foresight into pest

management is due to the fact that managing house flies offers a classic opportunity for IPM. An

effective house fly management strategy includes sanitation and other cultural methods,

mechanical control, insecticidal control, and, more recently, biological control (West 1951,

Kolbe 2004). Greene and Breisch (2002) documented the success of IPM in urban environments

by comparing the number of pest control service requests. From 1988 to 1999, the requests for

the group that included house flies were decreased to half its original number. Also, the amount

of insecticide used was drastically decreased to about 5% of its original use.

Sanitation is the primary means of managing house fly problems by eliminating breeding

sites and larval food, which also attract adult flies (Clymer 1973, Gojmerac 1979, Meyer and

Shultz 1990). Thus, by eliminating trash and manure, all stages of the fly life cycle are

disrupted.

In the urban setting, the main breeding site for the house fly is garbage. Kolbe (2004)

offers insight into the management of garbage for house fly control. Food in trash cans and

dumpsters should be picked up frequently. Twice a week is ideal. Trash cans and dumpsters

should have tight-fitting lids, and should be kept closed. The garbage placed in trash cans and

dumpsters should be sealed off in plastic bags to minimize attractive odors. Dumpsters should

be kept as far away from building entrances as possible. Also, keeping the dumpster and the

ground underneath it clean will help reduce house fly numbers.









In the agriculture, the main breeding site is manure (West 1951). Agricultural settings that

have accumulated manure have higher house fly populations (Lysyk and Axtell 1986). Feedlots

should allow for proper drainage that minimizes wet spots and prevents the build up of waste.

Manure and spilled feed should be removed twice a week (Clymer 1973). Sanitation, especially

the removal of manure, of poultry houses and dairies is also an essential tool in an integrated pest

management strategy against house flies (Lysyk and Axtell 1986, Lazarus et al. 1989). Manure

removal is most effective in all instances when it is done in the late spring (Axtell 1970a). Also

the bedding material that is routinely excreted upon by livestock may be changed or added to-

sawdust or sand-in order to make the manure less enticing as a breeding site for house flies

(MacCreary and Haenlein 1962, Schmidtmann 1988, Lazarus et al. 1989).

Another cultural method to prevent house fly problems is exclusion. House flies are

anthropomorphic and move to human structures through temperature and odor attractants. A

way to combat this invasion is to make sure that all windows and doors have screening, that the

screens lack holes, and are properly fitted (West 1951). House flies can also be excluded by

fans and air doors between rooms (West 1951, Hedges 1994).

Mechanical control is another method critical to an effective house fly management

regimen. Most likely the oldest control measure for house flies falls under this category -

flyswatting (West 1951). More modem methods of mechanical control are the plethora of

available fly traps. The different types of fly traps include baited traps, light traps, and sticky

traps (West 1951, Pickens et al. 1975, Skovmand and Mourier 1986, Pickens 1989, Kaufman et

al. 2005b). Of these types of traps there are a variety of different designs, attractants, and killing

methods. These are effective supplements to other control methods because they are able to trap

flies up to 24 hours a day, seven days a week with minimal effort (Kolbe 2004).









Insecticides can be used to treat a house fly problem. Chemicals have been used to control

flies since before World War II (West 1951). But, this should not be done indiscriminately, but

rather after other methods have failed to reduce the fly populations (Hedges 1993). This is due

to the house fly's ability to develop resistance to almost every insecticide used against it

(Learmount et al. 2002, Scott et al. 2000, Gao and Scott 2006). Not only does overuse of

insecticides contribute to house fly resistance, but there are also the concerns of public

perception, increased cost, safety, and environmental considerations (Carson 1962, Appling

1988, Stinson 1989). Different formulations should be considered for house fly management

based on the situation. They are residual insecticides applied to adult resting sites and larval

breeding sites, baits, and aerosols and ultra-low-volume (ULV) applications used as space

treatments (Kolbe 2004). Insecticide treatment should be used with caution as it may hinder a

proper IPM program i.e. affect natural predators that could be used as biological control agents

(Wills et al. 1990).

Augmentative biological control is the newest form of house fly control and can be used as

part of a successful IPM program. Fly egg predators represent one type of biological control, the

two major species being Carcinopspumilio (Erichson) and Machrocheles muscadomesticae

(Scopoli) (Axtell 1970b, Geden and Stoffolano 1987). Carcinopspumilio is capable of causing

-95% mortality of house flies, whereas M. muscadomesticae can cause -80% mortality in the

lab, with somewhat lower results in the field due to abiotic factors and the presence of alternate

prey (Geden and Axtell 1988, Geden et al. 1988). Fly larval predators also exist as biological

control agents. The black dump fly, Ophyra aenescens (Wiedemann), is a predator of house fly

larvae and is capable of out-competing the house fly (Nolan and Kissam 1985). Although O.

aenescens is a filth fly, it remains a suitable candidate for biological control because it is not a









nuisance. This is due to the adult habit of staying close to the breeding medium and its

preference for woods and shade as opposed to human structures (Legner and Dietrick 1974,

Nolan and Kissam 1987). Muscovy ducks are predators of adult house flies that may be used as

a biocontrol agent. The ducks ingest about 25 house flies per 15 minutes, and controlled 90% of

the house flies in a much shorter time than fly traps or baits (Glofcheskie and Surgeoner 1990).

Another group of biological control agents are the Pteromalid parasitoids that attack house

fly pupae. The most common are Muscidifurax raptor Girault and Sanders, Muscidifurax

zaraptor Kogan and Legner, Muscidifurax raptorellus Kogan and Legner, Spalangia cameroni

Perkins, Spalangia endius Walker, and Spalangia nigroaenea Curtis (Rueda and Axtell 1985,

Havron and Margalit 1991, Geden and Hogsette 2006). These are naturally occurring species

that hardly ever reach peak numbers and activity before or during the peak house fly nuisance

levels, therefore, the parasitoid populations must be augmented either by inoculative or

inundative release (Legner and Brydon 1966, Skovard and Nachman 2004). Skovard and

Nachman (2004) reported that the inundative release of S. cameroni usually caused house fly

populations to remain under nuisance levels (13-25 house flies per animal) on both dairy and pig

farms. Single species releases of Muscidifurax spp. and Spalangia spp. have had mixed results

(Geden et al. 1992, Geden and Hogsette 2006). Combined releases of the two genera are

preferred due to Muscidifurax spp. searching for hosts on the surface of manure, and Spalangia

spp. searching deeper (Rueda and Axtell 1985). Muscidifurax raptorellus and S. cameroni

simultaneously released accounted for up to -80% pupal mortality (Geden and Hogsette 2006).

Microbes can also provide biological control of the house fly. One example is the

entomopathogenic fungus, Entomophthora muscae Cohn. The fungus causes adult house fly

death in about 5-8 days after infection (Zurek et al. 2002). Once infected, house flies have a









100% mortality rate (Steinkraus and Kramer 1987). Infection of other individuals is likely due to

the males' preference to mate with infected females (Zurek et al. 2002). Another aspect that

makes E. muscae a good biological control agent is its ability to discharge conidiophores from

the cadaver of the dead fly (Zurek et al. 2002). A second entomopathogenic fungus that offers

promise is Beauveria bassiana (Balsamo). In laboratory studies, up to 98% mortality of house

flies after 15 days of exposure has been reported (Lecuona et al. 2005). In the field, B. bassiana

treated facilities had lower house fly populations than facilities treated with pyrethrins (Kaufman

et al. 2005a).

Nematodes in the genus Heterorhabditis, Steinernema, and Paraiotonchium have been

tested against house flies in the laboratory. While some show the ability to infect house flies and

show promise as control agents in the lab (Paraiotonchium muscadomesticae Coler caused 98%

mortality with a huge concentration of nematodes), the nematodes' infectivity is highly reduced

in manure, requiring larger application concentrations (Mullens et al. 1987, Geden 1997).

Fly Traps

There are three common types of house fly traps: light traps, sticky traps, and baited traps.

Light traps consist of an ultraviolet lamp that attracts the house fly into an electrocuting grid

(ELT) or a glue board (GLT) (Pickens 1994). Although ELTs are successful, GLTs are

becoming increasingly popular due to the noise emitted every time an insect is killed, and

because when electrocuted, house flies can scatter body parts and bacteria throughout the air

(Pickens 1989). Though Kolbe (2004) reports that low-voltage ELTs do not introduce fly

fragments into the air, Pickens (1989) observed a scatter of parts up to 1.5 m from the trap. Light

traps should be placed 0.3 to 0.9 m from the ground about 15 m apart around common fly

gathering places to maximize their effectiveness (Driggers 1971, Gilbert 1984, Pickens 1994).

Pickens et al. (1969) reported almost 100% capture of house flies in a 7.6 m2 room in only seven









hours. However, Thimijan et al. (1970) reported 0 to 16.3% catch of released house flies. It is

not surprising that some authors suggest that light traps are not sufficient control devices and

should only be used for monitoring (Schiefer 1970, Skovand and Mourier 1986, Rambo 1995).

Another type of trap that may be employed in an IPM program for house fly management

is sticky traps. One common type of sticky trap is a roll of paper with glue adhesive, usually

made attractive by color or printing patterns such as other flies on it. Kaufman et al. (2005b)

found that these traps caught over 80,000 house flies on each of five dairy calf greenhouse

facilities in only four days. They concluded that the sticky trap was responsible for a decline in

house fly populations and in each case, the producer noticed a reduction in house fly numbers

and was pleased. However, the efficacy of this type of trap declined in dusty environments, such

as occurs on many poultry farms (Kaufman et al. 2001). Another common type of sticky trap is

a pyramid sticky trap. Given the numbers of house flies trapped, these seem to be better suited

for monitoring instead of control. Pickens and Miller (1987) caught 5.6 to 14.0% of released

house flies with pyramid traps. They also point out that after about two days, the adhesive

became saturated with flies and was unable to catch significant numbers. Pickens et al. (1986)

and Johnson and Campbell (1987) describe pyramid traps as being most useful as monitoring

devices, not control devices, for Musca vetripennis (Say) and Musca autumnalis De Geer

respectively. Sticky tapes are also used as monitoring devices (Pickens et al. 1972, Turner Jr.

and Ruszler 1989).

Baited traps are probably the oldest of the fly traps, having been in use since about 1911

(Pickens 1995). The baited fly trap consists of an odiferous attractant inside of the trap that

entices the house flies to enter through an inverted cone (Bishopp and Henderson 1946, Pickens

1995). Similar to the other trap types, there is debate whether baited traps should be used as









monitoring or control devices. Dadour and Cook (1992) point out that to reduce a fly population

by 50-90%, a trapping regimen must remove 24-58% of the house flies. Pickens and Miller

(1987) report catching only 4.4 to 20.0% of released flies in baited traps. Smallegange (2004)

states that although baited traps show promise, they are not yet effective enough to reduce

populations to acceptable levels. Das (1994) reported that using Redtop Flycatcher baited with

ready made protein meal was easier and less expensive to use than a Baygon bait, but admitted

that it did not catch as many house flies as the Baygon bait killed. Burg and Axtell (1984) and

Geden (2005) also refer to baited traps as monitoring devices. Despite Dadour and Cook's

(1992) pessimism, Cohen et al. (1991) attained 64% reduction in house fly density, and

attributed a 42% reduction in diarrhea visits and 85% reduction in shigellosis visits of Israeli

soldiers to the trapping. Perhaps some of the low baited trapping numbers can be attributed to

how the baited traps are placed, after all, daily catches of up 40,000 to 129,000 have been

reported (Blair 1945, Pickens 1995). Also, seasonal catches of 1.1 to 9.7 million have been

recorded (Fenton and Bieberdorf 1936, Mer and Paz 1960). Baited traps should be placed

outside every 6-9 m and placed as close as possible to common fly-breeding materials (Pickens

1994). Traps are most effective when placed within 0.3 m of the ground (Mitchell and Tingle

1975).

Objectives

The first objective of these experiments was to discover the relative effectiveness of each

of six commercial fly trap designs in capturing house flies. The second objective was to

determine the attractiveness of yeast and ammonium carbonate to house flies, and whether

combining and/or aging these ingredients would affect their attractiveness. Using this

information, I would be able to make a few general conclusions as to where commercial fly traps

and the tested attractants fit into an IPM program for house flies.









CHAPTER 2
MATERIALS AND METHODS

Laboratory Rearing of the House Fly

House fly larvae, M. domestic, Horse Teaching Unit (HTU) strain established in 2004,

from Gainesville, FL, were reared in plastic tubs (33.50 by 28.41 cm) on medium containing 250

ml of Calf Manna (Manna Pro Corp., St. Louis, MO), 1.5 liters of water, and 3 liters of wheat

bran. The pupae were separated from the medium by water floatation on the seventh day and

dried on paper toweling. The pupae were placed in screened rearing cages (40.64 by 26.67 by

26.67 cm) for emergence. The screened rearing cages were screened on five sides with the sixth

side used as an entryway that was covered with surgical stocking net that could be tied off to

prevent house fly escape. Adult house flies were provided granulated sugar, powdered milk, and

water ad libitum (Hogsette et al. 2002). Both larvae and adults were held on a 12:12 (L:D)

photoperiod, 26.2 + 0.50 C, and 51.2 + 3.5% RH. Adults emerged in the cage following three

days as pupae. Oviposition was induced with spent larval medium at around ten days post

eclosion. The eggs collected were used to sustain the colony. Males and females were held

together in the same cages.

Adult house flies were aspirated from screen cages with a modified hand-held vacuum.

Aspirated house flies were placed into a freezer (-300 C) until inactive (-1-5 min). After

removal from the freezer, the house flies were placed on a chilled aluminum tray. House flies

were counted and sexed (25 males, 25 females) and held in a deli cup (236.58 ml) for about one

hour. House flies used in the design experiment and the aged attractant combination test were

between three and seven days post-eclosion. House flies used in scatter and attractant

component experiments were between three and five days post-eclosion.









Fly Trap Design Experiments


Scatter Experiment

An open release cage study was developed to evaluate the house flies' propensity to escape

from an enclosed space in a 24 hour period. Two different trap designs were utilized no trap,

designated the laboratory room, and a CDC (Center for Disease Control) net trap (Fig. 2-1).

Each release cage (28.8 by 26.1 by 39.1 cm high) consisted of a screen cage fitted with a

stocking net opening. A cotton-filled plastic cup (88.72 ml) soaked with 50 ml of 10% aqueous

sugar solution was placed in release cage. House flies were then released from the deli cup into

each release cage. A continuous channel was created by fitting a yellow funnel that was

removed from a Trap n' TossTM (Farnam Companies, Inc., Phoenix, AZ) fly trap. The stocking

net opening of the release cage was made continuous with the cone. One half of the release

cages were oriented so that the cone opening was below the cage, while the other half were made

so that the cone opening was above the release cage (Fig. 2-2 and 2-3). For the release cages that

were left open to the laboratory room, after 24 h, the sugar water was removed, the cone

removed, and the release cage was tied off for 24 h to kill the house flies remaining in the release

cage. The dead house flies were counted and recorded to determine the percentage of house flies

that escaped into the laboratory room by taking the difference of how many house flies were left

in the release cage as opposed to how many were initially released. For those release cages fitted

with CDC net traps, the CDC net traps were taken from the release cage, tied off and left to sit

for 24 h to kill the flies. The flies were then counted, sexed, and recorded to determine the

percentage of male, female, and total house flies that were trapped in the CDC net traps. A

replication consisted of a cage for top-entry or bottom-entry treatments. Four replications were

of laboratory room catch at the same time in the laboratory (22.8 0.02 C) on a 24:0 (L:D)

photoperiod. The same procedure was done for the experiments with CDC net traps.









Commercial Fly Trap Experiment

Fly traps

A comparison of six commercial fly traps was conducted in the laboratory. Fly traps used

were categorized as either bottom-entry or top-entry. Bottom-entry fly traps were designed with

an inverted funnel leading into a collection container. Bottom-entry fly traps included Trap n'

TossTM, The Advantage Fly TrapTM (Advantage Traps, Inc., Columbia, SC), and BC 1752 Dome

(McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK) (Figs. 2-4, 2-5, and 2-6).

Top-entry fly traps were containers fitted with entrance(s) on the lid. Rescue! Reusable

Fly Trap (Sterling International, Inc., Liberty Lake, WA) and Victor Fly Magnet Trap

(Woodstream, Lititz, PA) each had four entrance holes, while the Fly Terminator Pro (Farnam

Companies, Inc., Phoenix, AZ) had only one entrance hole (Figs. 2-7, 2-8, and 2-9).

The diameter for each entrance/exit hole were measured and used to calculate area. The

areas were used to calculate the entrance:exit ratios for each fly trap. The slope of the funnel

entrance and the volume of each fly trap's capacity were also measured. Each entrance color

was observed and recorded.

Bioassay

A fly attractant mixture consisting of 5 g yeast, 0.12 g ammonium carbonate, and 75 ml of

water was placed into each fly trap immediately after mixing. A cotton-filled plastic cup soaked

with 50 ml of 10% aqueous sugar solution was placed in release cage. House flies were released

from the deli cup into release cage. The stocking net opening of the release cage was made

continuous with the opening of the fly trap, which were either placed on top of the release cage

(bottom-entry fly traps) or suspended below the release cage (top-entry fly traps) (Figs. 2-10 and

2-11). The stocking net was pulled over the set fly trap and tied off to prevent house flies from

escaping between the release cage and the fly trap.









After 24 h, each fly trap was removed from the release cage, placed into a sealable plastic

bag, then refrigerated at 30 C for 24 h. Upon removal from the refrigerator, captured house

flies were extracted from the attractant mixture using a colander, placed on a chilled aluminum

tray, counted, and sexed.

The experiment was a completely randomized design comparing each fly trap's capture

efficacy. Each trial was run in the laboratory (22.8 0.020 C) on a 24:0 (L:D) photoperiod. The

locations of the fly traps were randomized to minimize position effect. Testing included one of

each fly trap (bottom-entry and top-entry) examined five times, except Fly Terminator Pro

which was left out once due to availability. On two additional occasions, bottom-entry traps

were tested using two Trap n' TossTM fly traps, one Advantage Fly TrapTM, and one BC 1752

Dome Trap.

Attractant Experiments

Aged Attractant Experiment

The Trap n' TossTM fly trap was used as a standardized fly trap to evaluate the effect of

attractant age on fly capture. Sets of three groups of fly attractants were prepared separately

using 5 g yeast, 0.12 g ammonium carbonate, and 75 ml of water. These mixtures were held for

0, 1, 2, 3, or 4 d at 22.8 0.020 C yielding five age classes of attractant. Following the aging of

each attractant, the attractants were introduced into individual fly traps. A cotton-filled plastic

cup containing 10% aqueous sugar solution and the 50 house flies were introduced as previously

described. The fly trap was secured on the cage, as before. Following the 24 h experiment,

house flies were collected, counted, and sexed as in commercial trap experiments. Three cages

of each attractant age mixture were run simultaneously (22.8 0.020 C) on a 24:0 (L:D)

photoperiod. The experiment was run twice. Due to contamination of release cage with

insecticide, one cage of 1-d aged attractant was discarded.









Attractant Component Experiment

This study consisted of four treatments including 75 ml water control, 0.12 g ammonium

carbonate, 5 g of yeast, or a combination of the ammonium carbonate and yeast, each in 75 ml of

water. Each of the attractants was aged 0, 2, or 4 d prior to placement into a Trap n' TossTM fly

trap. A cotton-filled plastic cup containing 10% aqueous sugar solution and the 50 house flies

were introduced, then the stocking net used to secure the fly trap on the cage, as before. House

flies were collected, counted, and sexed as in commercial trap experiments. Each combination

was run simultaneously and was run independently six times, for a total of six replications.

Experiment was run in the lab (22.8 + 0.02 C) on a 24:0 (L:D) photoperiod.

Data Analysis

Actual means are presented in tables. Data for percentage catch of males, females, and

total for laboratory room, CDC net trap, each commercial trap, and each aged attractant were

arcsine square root transformed and analyzed using one-way analysis of variance. Male, female,

and total percent house fly catch was analyzed by linear regression for the attractant mixture

aged 1-4 days. A one-way analysis of variance was run on each attractant component by days

aged (0, 2, 4). Means for total, male, and female percent catch were separated using Student

Newman-Keuls (SNK) Test (P = 0.05) (SAS 2001). A Student's t-test (P = 0.05) was used to

compare the entrance position variable for laboratory room versus CDC net trap, the catch

variable for laboratory room versus CDC net trap catch, and the sex variable for the different

ages and combinations of attractants.








































Figure 2-1. CDC Net Trap used to capture house flies escaping from screen cages.




































Figure 2-2. Bottom-entry release cage design fitted with a Trap n Toss' cone and open to
laboratory room.




































Figure 2-3. Top-entry release cage design fitted with modified Trap n Toss' and open to
laboratory room






































Figure 2-4. Trap n' Toss'" (Farnam Companies, Inc., Phoenix, AZ ) fly trap



































Figure 2-5. The Advantage Fly TrapTM (Advantage Traps, Inc., Columbia, SC)


































Figure 2-6. BC 1752 Dome (McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK)




































Figure 2-7. Rescue! Reusable Fly Trap (Sterling International, Inc., Liberty Lake, WA)











































Figure 2-8. Victor Fly Magnet Trap (Woodstream, Lititz, PA)











































Figure 2-9. Fly Terminatoi


Pro (Farnam Companies, Inc., Phoenix, AZ)





































gure 2-10. Bottom-entry study design with Trap n' Toss'" (Farnam Companies, Inc.,
AZ) fly trap attached.


loenix,




































gure 2-11. Top-entry study design w
attached.


Victor Fly Magnet Trap (Woodstream, Lititz, PA)









CHAPTER 3
RESULTS

Fly Trap Design Experiments

Scatter Experiment

When the house flies were released from the deli cup into the release cage, they exhibited

resting behavior preceded by a short time of flying or walking. They did not show any initial

observational attraction to the trap, sugar water, or opening of the release cage. However, over

half (53.38 + 4.18%) were eventually captured. The cone allowed house flies to easily leave the

release cage, but made it difficult to reenter.

The percentage of house flies both captured in CDC net traps and escaped into the

laboratory room after 24 h was determined (Table 3-1). For the released male house flies, there

was no significant difference between bottom-entry and top-entry treatments (df = 9.11, t = 2.08,

P = 0.0669). There was no significant difference in house fly escape between bottom-entry

(82.00 6.63%) and top-entry (90.00 2.58) treatments for the laboratory room. However,

significantly more males escaped into the laboratory room than escaped into CDC net traps (df=

9.34, t= 4.39, P = 0.0016). Significantly more males escaped through top-entry (67.00 2.52%)

treatments than into the bottom-entry (22.00 3.46%) treatments into CDC net traps (df = 3, F =

22.13, P < 0.0001). Within the bottom-entry treatment, significantly more males escaped into

the laboratory room (82.00 6.63%) than were caught in the CDC net trap (22.00 3.46%)

(Table 3-1). Significantly more male house flies escaped into the laboratory room (90.00 +

2.58%) than into the CDC net trap (67.00 2.52%) within the top entry treatment.

For the released female house flies, there was no significant difference between bottom-

entry and top-entry treatments (df = 14, t = 0.20, P = 0.8457). Regardless of treatment,

significantly more female house flies escaped into the laboratory room than were captured in









CDC net traps (df= 8.15, t = 6.43, P = 0.0002). There was no significant difference in house fly

escape between bottom-entry (67.00 + 11.12%) and top-entry (65.00 11.24%) treatments for

the laboratory room escape option. Overall, few female house flies were captured in the CDC

net traps (17.00 2.97%) and there was no significant difference between bottom-entry (19.00 +

2.52%) and top-entry (15.00 + 3.42%) treatments. Within the bottom-entry treatment,

significantly more females escaped into the laboratory room (67.00 + 11.12%) than were

captured in the CDC net trap (19.00 2.52%). Significantly more female house flies also

escaped into the laboratory room (65.00 11.24%) than into the CDC net trap (15.00 3.42%)

for the top-entry treatment (df = 3, F = 11.50, P = 0.0008).

There was no significant difference between the total numbers of house flies that escaped

through top entry and bottom entry treatments (df = 14, t = 0.90, P = 0.3844). Overall, house

flies were not caught in CDC net traps as often as they escaped into the laboratory room (df = 14,

t = 7.67, P < 0.0001). About three quarters of the released house flies escaped into the laboratory

room (76.00 4.04%). Top-entry CDC net traps (41.00 2.89%) caught significantly more

house flies than the bottom-entry CDC net traps (20.50 2.87%) (df = 3, F = 27.49, P < 0.0001).

There was no significant difference between bottom-entry (74.50 6.99 %) and top-entry (77.50

+ 5.06%) traps. The mean trap catch for the CDC net traps was only one-third (30.75 4.31%)

of the total released house flies. However, significantly more house flies escaped through

bottom-entry traps into laboratory room (74.50 6.99%) than were captured in the bottom-entry

CDC net trap (20.50 2.87%) (df= 3, F = 27.49, P < 0.0001). This pattern was also observed

with top-entry traps where significantly more house flies escaped into the laboratory room (77.50

5.06%) than into the CDC net trap (41.00 2.89%) (df = 3, F = 27.49, P < 0.0001).









Commercial Fly Trap Dimensions

Six commercial fly traps were compared for their ability to capture house flies released

into screened cages (Table 3-2). Each fly trap consisted of a plastic container that held both the

attractant and the captured house flies. All fly traps had funnels that had both entrance and exit

holes that facilitated house fly capture but limited escape. Every fly trap, except the

Terminator Pro, had entrance holes that were larger than the exit holes. The Trap n' TossT, a

bottom-entry trap, had the greatest entrance area of all the fly traps (176.24 cm2), whereas the

Fly Terminator Pro had the largest entrance area for the top-entry fly traps (20.51 cm2). The

smallest entrance area of the top-entry fly traps was the Rescue! Reusable Fly Trap (3.16 cm2),

whereas the smallest bottom-entry area was The AdvantageTM Fly Trap (46.57 cm2). The

smallest exit area of the top-entry fly traps was the Rescue! Reusable Fly Trap (2.00 cm2),

whereas the bottom-entry fly trap with the smallest exit area was The Advantage Fly TrapT

(9.08 cm2). The largest exit area of the top-entry fly traps was the Fly Terminator Pro (20.51

cm2), whereas the Trap n' TossTM had the largest exit area (11.95 cm2) of bottom-entry traps.

The fly trap entrance to exit ratio is a measure of the entrance area divided by the exit area.

This is an important measurement because it provides the best relative combination of allowing

house flies into the fly trap without releasing them. The bottom-entry Trap n' TossTM had the

greatest entrance:exit ratio of the fly traps (14.75), whereas the Fly Magnet had the greatest

entrance:exit ratio of the top-entry fly traps (2.05). The top-entry Fly Terminator Pro had the

smallest entrance:exit ratio of all the fly traps (1.00), whereas BC 1752 Dome had the smallest

entrance:exit ratio of the bottom-entry fly traps (4.64).

The slope of the entrance cone directs the house flies into the holding container and the

range of slopes observed by these traps was considerable. The BC 1752 Dome had the greatest

slope of the bottom-entry fly traps (75.570), whereas the Terminator Pro, a top-entry fly trap,









had the greatest slope of all the fly traps (90.000). The Trap n' TossTM, a bottom-entry fly trap,

had the least slope of all the fly traps (61.670), whereas the Rescue! had the least slope of the

top-entry fly traps (85.070).

The bottom-entry fly trap with the largest fly holding volume was Trap n' TossTM (1936

ml), whereas the fly trap with the largest volume was the top-entry Fly Terminator Pro (3786

ml). The fly trap with the smallest volume was the BC 1752 Dome, whereas the top-entry fly

trap with the smallest volume was the Fly Magnet (983 ml).

Each fly trap's entrance was one of several colors. The Trap n' TossTM and BC 1752

Dome's entrances were colored yellow. The Advantage Fly TrapTM's entrance was white. The

Fly Magnet and Fly Terminator Pro had black entrances, while the Rescue! had a green

entrance.

Commercial Fly Trap Experiment

When house flies were released from deli cups into the release cage, they showed the

previously described initial behavior. At the completion of the 24 h experiment, some of the

house flies were attracted to or wandered into the fly traps. However, it is likely that some may

have escaped back into the release cage. House flies would continue to show this basic behavior

throughout the experiments.

The percentage of house flies captured in six different commercial fly traps after 24 h was

recorded (Table 3-3). For this experiment, the mean catch of released males was 36.67 3.17%

(n = 42). The highest capture was with the The Advantage Fly TrapTM (51.43 7.33%), while

the lowest catch was with the Fly Terminator Pro (22.00 13.71%). These were the only

significantly different catch means with respect to released male house flies (df = 5, F = 2.87, P =

0.0303). There was no significantly superior bottom-entry fly trap when compared only to the

other bottom-entry traps or top-entry fly trap when compared only to other top-entry traps.









The mean catch of released females for this experiment was 41.90 3.56% (n = 42). The

Trap n' TossT caught significantly more released females (66.22 5.89%) than the Advantage

(35.43 6.32%), BC 1752 Dome (34.86 6.97%), Fly Magnet (32.80 7.31%), and Fly

Terminator Pro (28.00 5.42%) (df = 5, F = 4.58, P = 0.0031). Rescue! (58.40 12.24%)

was not significantly different than the best or the worst fly traps when catching female house

flies. There was no significant difference between the top-entry fly traps; however, the Trap n'

TossTM caught significantly more house flies than the other two bottom-entry fly traps (df = 5, F

= 4.58, P = 0.0031).

The mean capture for all fly traps in the experiment was 39.33 2.58% of the total

released house flies (n = 42). The fly trap that captured the most house flies was the Trap n'

TossT (52.22 5.13%), while the Fly Terminator Pro captured the fewest (25.00 + 8.10%).

These were the only significantly different catch means with respect to total house flies released

(df = 5, F = 3.46, P = 0.0134). There were no significantly superior fly traps when comparing

only top-entry fly traps with each other and bottom-entry fly traps with each other.

Attractant Experiments

Aged Attractant Experiment

The percentage of house flies after captured after 24 h in fly traps with a yeast and

ammonium carbonate mixture aged 0 4 days (by one day intervals) was determined (Table 3-4).

A significant difference between aged attractants was observed with male house flies (df= 4, F =

3.44, P = 0.0233). Attractants aged for four days (62.00 5.54%) and three days (60.67 +

7.04%) caught significantly more house flies than 1-day aged attractant (35.20 5.43%). The

freshly mixed attractant (38.22 3.49%) and 2-day aged attractant (42.00 7.85%) were not

significantly different from the mixtures that captured the most and least house flies. Regression









analysis demonstrated that as attractant aged from one to four days, male house fly catch

increased linearly (y = 9.907x + 25.2, R2 = 0.9073) (Fig. 3-1).

As with males, a significant difference was also observed between female house fly

capture among the attractants (df = 4, F = 3.63, P = 0.0188). The 4-day aged attractant (78.00 +

5.91%) caught significantly more females than attractant aged 1 day (47.20 4.80%).

Significant differences were not observed among other treatments. Regression analysis

demonstrated that as attractant aged from 1 to 4 days, female house fly catch increased linearly

(y = 11.173x + 31.2, R2 = 0.8801) (Fig. 3-1).

Overall, there was a significant difference between the numbers of house flies captured

using variously aged attractants (df = 4, F = 4.62, P = 0.0065). The 4 day aged attractant (70.00

4.73%) captured significantly more house flies than attractants aged two days (49.00 6.65%)

and one day (41.20 4.08%). Attractant aged three days (63.00 5.99%) caught significantly

more house flies than attractant aged one day (41.20 4.08%). Freshly mixed attractant (55.33 +

3.13%) was not significantly different from any of the other mixtures. Regression analysis

demonstrated that as attractant aged from 1 to 4 days, total house fly catch increased linearly (y =

10.04x + 30.7, R2 = 0.9827) (Fig. 3-1).

Attractant Component Experiment

To determine the importance of attractant age and attractant components on house fly

capture, a study was conducted examining individual and combinations of components at 0, 2,

and 4 days post-mixing (Table 3-5). The freshly mixed combination treatment (56.00 7.93%)

captured significantly more male house flies than the ammonium carbonate (20.00 6.20%) and

the control treatments (9.33 3.04%) (df = 3, F = 9.31, P = 0.0005). For attractants aged both 2

and 4 days, the combination and yeast treatments captured significantly more house flies than the









ammonium carbonate and control treatments, respectively (df = 3, F = 10.94, P = 0.0002; df= 3,

F = 8.45, P = 0.0008).

The freshly mixed and 4-day-old attractant, combination and yeast treatments captured

significantly more female house flies than the ammonium carbonate and control treatments,

respectively (df = 3, F = 12.97, P < 0.0001; df = 3, F = 16.98, P < 0.0001). For attractants aged 2

days, the combination treatment (69.33 8.56%) caught significantly more house flies than the

yeast treatment (40.67 8.48%), which caught significantly more house flies than ammonium

carbonate (18.67 6.25%) and the control treatments (12.67 7.26%) (df = 3, F = 10.21, P =

0.0003).

The freshly mixed combination treatment (61.00 5.48%) captured significantly more

total house flies than the yeast treatment (41.67 5.55%), which captured significantly more

house flies than the ammonium carbonate (22.00 6.85%) and the control treatments (9.67 +

3.32%). With both 2-day and 4-day aged attractants, the combination and yeast treatments

captured significantly more house flies than the ammonium carbonate and control treatments,

respectively (df = 3, F = 11.77, P = 0.0001; df= 3, 24.27, P < 0.0001).

The combination (df = 10, t = 2.91, P = 0.0155) and yeast (df = 10, t = 2.56, P = 0.0283)

treatments captured significantly more female house flies than males when aged for 4 days. All

other combinations of attractant components and ages did not show significant differences

between the sexes.









Table 3-1. Percent escape of male, female, and total house flies into CDC net traps or laboratory room.
% (SE)a


Trapb


n Males


Females


Total


Bottom Entry
Room
Net

Top Entry
Room
Net


82.00 (6.63)a
22.00 (3.46)c


90.00 (2.58)a
67.00 (2.52)b
df = 3, F = 22.13, P < 0.0001


67.00 (11.12)a
19.00 (2.52)b


65.00 (11.24)a
15.00 (3.42)b
df= 3, F = 11.50, P = 0.0008


74.50 (6.99)a
20.50 (2.87)c


77.50 (5.06)a
41.00 (2.89)b
df = 3, F = 27.49, P < 0.0001


SData were arcsine square root transformed before analysis. Means within a column followed by the same letter are not significantly
different (c = 0.05, Student Newman-Keuls test; SAS 2001).
a 25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Fly trap removed
from release cage following 24 h exposure.
b Room = house flies allowed to escape into laboratory room. Net = house flies captured in CDC net trap affixed to cone opening.
C n = replications










Table 3-2. Characteristics of six commercially available fly traps used to evaluate house fly capture efficacy.


Trapa


Entrance (cm2)


TM
Trap n' Toss


AdvantageTM

BC 1752 Dome

Rescue!

Fly Magnet

Terminator Pro


176.24


46.57

49.00

3.16

6.16

20.51


Exit (cm2)


11.95


9.08

10.56

2.00

3.00

20.51


Ratiob

14.75


5.13

4.64

1.58

2.05

1.00


Slope (deg)


61.67


75.33

75.57

85.07

89.97

90.00


Volume (ml)c


1936


1840


1060


3786


a Bottom-entry traps were Trap n' TossTM (Farnam Companies, Inc., Phoenix, AZ), The Advantage Fly TrapTM (Advantage Traps,
Inc., Columbia, SC), and BC 1752 Dome McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK), and were placed on top of the
release cage. Top-entry traps were Rescue! Reusable Fly Trap (Sterling International, Inc., Liberty Lake, WA), Victor Fly Magnet
Trap (Woodstream, Lititz, PA), and Fly Terminator Pro (Farnam Companies, Inc., Phoenix, AZ), and were placed under the release
cage.
b Ratio = entrance + exit
' Volume = number of ml holding capacity of fly trap


Color

yellow


white

yellow

green

black

black









Table 3-3. Percentage of male, female, and total house flies caught in six commercial traps.


Trapb


Males


Trap n' TossT

AdvantageTM

BC 1752 Dome

Rescue!

Fly Magnet

Terminator Pro


38.22 (5.21)ab

51.43 (7.33)a

22.86 (4.07)ab

36.00 (9.38)ab

24.80 (5.57)ab

22.00 (13.71)b


% Catch (SE)a


Females


66.22 (5.89)a

35.43 (6.32)b

34.86 (6.97)b

58.40 (12.24)ab


Total


52.22 (5.13)a

43.71 (4.40)ab

28.86 (5.03)ab

47.20 (9.73)ab

28.80 (6.09)ab

25.00 (8.10)b


32.80 (7.31)b

28.00 (5.42)b


df = 5, F = 2.87, P = 0.0303


df = 5, F = 4.58, P = 0.0031


df = 5, F = 3.46, P = 0.0134


Data were arcsine square root transformed before analysis. Means within a column followed by the same letter are not significantly
different (c = 0.05, Student Newman-Keuls test; SAS 2001).
a 25 male and 25 female, 3-7 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Fly trap removed
from release cage following 24 h exposure. All traps contained a combination of yeast and ammonium carbonate (41.67 : 1) in 75 ml
of water.
b Bottom-entry traps were Trap n' TossTM (Farnam Companies, Inc., Phoenix, AZ), The Advantage Fly TrapTM (Advantage Traps, nc.,
Columbia, SC), and BC 1752 Dome McPhail) Trap (AgrisenceTM Agriculture, Pontypridd, UK), and were placed on top of the release
cage. Top-entry traps were Rescue! Reusable Fly Trap (Sterling International, Inc., Liberty Lake, WA), Victor Fly Magnet Trap
(Woodstream, Lititz, PA), and Fly Terminator Pro (Farnam Companies, Inc., Phoenix, AZ), and were placed under the release cage









Table 3-4. Percentage of male, female, and total house flies captured after 24 h in Trap n' TossTM fly traps baited with an attractant
aged up to 4 d.


% Catch (SE)a


Attractant age (d)b


Males


Females


Total


6 38.22 (3.49)ab

5 35.20 (5.43)b

6 42.00 (7.85)ab

6 60.67 (7.04)a

6 62.00 (5.54)a

df = 4, F = 3.44, P


0.0233


59.33 (4.78)ab

47.20 (4.80)b

56.00 (7.00)ab

65.33 (7.06)ab

78.00 (5.91)a

df = 4, F = 3.63, P


0.0188


55.33 (3.13)abc

41.20 (4.08)c

49.00 (6.65)bc

63.00 (5.99)ab

70.00 (4.73)a

df = 4, F = 4.62, P


Data were arcsine square root transformed before analysis. Means within a column followed by the same letter are not significantly
different (c = 0.05, Student Newman-Keuls test; SAS 2001).
a25 male and 25 female, 3-7 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Trap n' TossTM
(Farnam Companies, Inc., Phoenix, AZ) fly trap removed from release cage following 24 h exposure.
b Combination of yeast and ammonium carbonate (41.67 : 1) in 75 ml of water
SReplications


0.0065









Table 3-5. Percentage of house flies caught in baited Trap n' TossTM fly traps with attractants aged up to 4 d.


Male % Catch (SE) a


Days


Attractantb


Control

Yeast


Ammonium
Carbonate

Combination


9.33 (3.04)c

34.00 (6.26)ab

20.00 (6.20)bc


56.00 (7.93)a

df= 3, F = 9.31, P


7.33 (2.40)b

45.33 (6.98)a

16.67 (5.97)b


46.00 (7.14)a


0.0005


df= 3, F = 10.94, P


10.67 (3.82)b

37.33 (5.81)a

17.33 (4.92)b


44.00 (7.08)a


0.0002


df = 3, F = 8.45, P = 0.0008


Data were arcsine square root transformed before analysis. Means within a column followed by the same letter are not significantly
different (c = 0.05, Student Newman-Keuls test; SAS 2001). n = 6
a25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Trap n' TossTM
(Famam Companies, Inc., Phoenix, AZ) fly trap removed from release cage following 24 h exposure.
bControl = 75 ml tap water, yeast = dried active baker's yeast in 75 ml of tap water, ammonium carbonate = ammonium carbonate in
75 ml of tap water, combination = yeast and ammonium carbonate (41.67 : 1) in 75 ml of tap water









Table 3-5 Continued.


Female % Catch (SE) a

Days


Attractantb


Control


Yeast


Ammonium
Carbonate

Combination


10.00 (4.70)b

49.33 (6.67)a

24.00 (8.20)b


66.00 (5.03)a


12.67 (7.26)c

40.67 (8.48)b

18.67 (6.25)c


69.33 (8.56)a


15.33 (3.78)b

61.33 (7.35)a

24.67 (5.79)b


77.33 (8.98)a


df= 3, F = 12.97, P < 0.0001


df= 3, F = 10.21, P = 0.0003


df= 3, F = 16.98, P < 0.0001


Data were arcsine square root transformed before analysis. Means within a column followed by the same letter are not significantly
different (c = 0.05, Student Newman-Keuls test; SAS 2001). n = 6
a25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Trap n' TossTM
(Farnam Companies, Inc., Phoenix, AZ) fly trap removed from release cage following 24 h exposure.
b Control = 75 ml tap water, yeast = dried active baker's yeast in 75 ml of tap water, ammonium carbonate = ammonium carbonate in
75 ml of tap water, combination = yeast and ammonium carbonate (41.67 : 1) in 75 ml of tap water









Table 3-5 Continued


Total % Catch (SE)a

Days


Attractantb


Control


Yeast


Ammonium
Carbonate

Combination


9.67 (3.32)b

41.67 (5.55)a

22.00 (6.85)b


61.00 (5.48)a


10.00 (4.41)b

43.00 (7.30)a

17.67 (5.85)b


57.67 (6.52)a


13.00 (3.09)b

49.33 (5.77)a

17.67 (3.56)b


60.67 (5.90)a


df= 3, F = 13.15, P < 0.0001


df= 3, F = 11.77, P = 0.0001


df = 3, F = 24.27, P < 0.0001


Data were arcsine square root transformed before analysis. Means within a column followed by the same letter are not significantly
different (c = 0.05, Student Newman-Keuls test; SAS 2001). n = 6
a 25 male and 25 female, 3-5 d post-eclosion, house flies released into 28.8 by 26.1 by 39.1 high cm release cage. Trap n' TossTM
(Farnam Companies, Inc., Phoenix, AZ) fly trap removed from release cage following 24 h exposure.
b Control = 75 ml tap water, yeast = dried active baker's yeast in 75 ml of tap water, ammonium carbonate = ammonium carbonate in
75 ml of tap water, combination = yeast and ammonium carbonate (41.67 : 1) in 75 ml of tap water













fen-male,











males
= 1 04. 5:
P- = O 9073


* Males
* Females
Total


Attractant Age

Figure 3-1. Effect of attractant age on percent catch of male, female, and total house flies.









CHAPTER 4
DISCUSSION AND CONCLUSIONS

Fly Trap Design Experiments

House flies tend to scatter from sites where they are released (Hindle 1914, Jones et al.

1999). Sacca (1963) concluded that house flies disperse in a more or less random pattern until

suitable conditions are found. However, Bishopp and Laake (1919) found that house flies would

pass up favorable feeding and breeding places in the course of dispersal. In my experiments, the

house flies released into a cage scattered from the cage into the laboratory room and various fly

traps. This scattering behavior was in accordance with the past literature. The house flies

scattered from the release cage, which contained nutrition, water, and potential mates. Sacca's

(1963) observation that house flies seem to move from one place to another in a random fashion

was true of the observations in my experiments. However, Sacca (1963) stated that these

random movement episodes end when suitable conditions are found i.e. food and mates, but the

behavior of the house flies in my experiments to forgo suitable conditions contradicted this.

Instead the house flies in my experiments behaved more in accordance with the findings of

Bishopp and Laake (1919). This behavior may account for why more house flies escaped from

the release cage into the laboratory room than the CDC net trap and the un-baited Trap n'

TossTM, because there is a small chance that the house flies would move back towards the

opening of the release cage and go back in, but as the house flies scatter into one of the fly traps,

they can just as well scatter back into the release cage.

However, despite the seemingly random scatter from the release cages, air movement may

also affect the house fly dispersal behavior. Several authors have found house flies to be

positively anemotropic (Hindle 1914, Hindle and Merriman 1914, Ball 1918, Carment 1922,

Uvarov 1931). In my experiments, more house flies scattered into the laboratory room than









ended up in the CDC net trap, and more were in the CDC net trap than in the un-baited Trap n'

TossTM. There is a direct correlation with allowed air movement in this situation. The bottom-

entry CDC net trap versus the top-entry CDC net trap difference in house fly catch suggest the

position of the trap was very important to return dispersal. For each treatment, there was the

same entrance/exit cone from the Trap n' TossTM, but the Trap n' TossT trap allowed less air

flow than did the CDC net trap, and the CDC net trap allowed less air flow than did the open

laboratory room. The behavior of the house flies in my experiments was in accordance with

previous authors' findings, that house flies will seek and fly upwind.

Most of the commercial fly traps were not significantly different in their abilities to capture

house flies. However, the Trap n' TossT caught significantly more house flies than the

Terminator Pro, and this was probably due to a number of design factors including large

entrance hole area, small exit hole area, the ratio of entrance to exit hole ratio, the slope being

close to 600, and the color of the entrance. This is not surprising when considering the overall

design of each fly trap. Pickens (1995) stated that the larger the entrance hole, the more house

flies that would be captured with a fly trap. His optimum was an entrance of 506.45 cm2. I

found the Trap n' TossT to be only -35% of that area, but the Fly Terminator Pro was a mere

4% of Pickens' (1995) ideal entrance area. So, it can be said that a large entrance may be

partially responsible for the Trap n' TossTM having a higher percent catch of the house flies than

Terminator Pro. Pickens (1995) also states that a small exit will work best to keep captured

house flies in the fly trap. His ideal exit area is 0.28 to 1.33 cm2. I found the Trap n' TossTM to

be almost nine times larger than 1.33 cm2, but Fly Terminator Pro is over 15 times larger. The

smaller exit area of the Trap n' TossTM may have contributed to its higher house fly capture

through retention of captured house flies. If a large entrance and a small exit are best for









capturing house flies, then, within reason, the higher the ratio of entrance to exit, the more house

flies a fly trap should catch. Again the Trap n' TossTM ratio was almost 15 times larger than that

of the Terminator Pro. The slope may have been a factor in the Trap n' TossT capturing more

house flies than Terminator Pro. According to Pickens (1995) the slope of the cone of the fly

trap should be 600. The Trap n' TossTM had a slope barely above 600, while Fly Terminator

Pro was 90.

There have been many studies documenting which color makes a surface most attractive to

house flies, and there have been varied conclusions. Awati and Swaminath (1920) used colored

paper and found yellow to be most attractive. Burg and Axtell (1984) painted plastic jug traps

and found yellow to be the most attractive to house flies. Pickens (1995) stated that white or

yellow were preferred in cool weather, while red, black and blue were the preferred color

surfaces when the weather was hot, while Muniz (1967) found that color preference was

independent of temperature. Hecht et al. (1968) using colored pieces of cardboard found that

black was most attractive in the laboratory, while yellow was most attractive outdoors. Nava

(1967) found that house flies were attracted to black cardboard more than any other color. The

studies of Awati and Swaminath (1920), Burg and Axtell (1984), and to some extent Hecht et al.

(1968) and Pickens (1995) substantiate that the yellow color may have played a role in the

yellow entrance of Trap n' TossTM capturing more house flies than the black entrance of Fly

Terminator Pro.

The commercial fly traps had many different characteristics, but only two fly traps were

significantly different from each other. The Trap n' TossT entrance area was almost nine times

larger and the exit area was about half as large as the Fly Terminator Pro. The ratio then of

entrance area to exit area for the Trap n' TossTM was almost 15 times more than Terminator









Pro. The slope of the entrance of Trap n' TossT was nearly the ideal 600, while Fly

Terminator Pro was -300 steeper. The color of the Trap n' TossT entrance was yellow,

whereas the Fly Terminator Pro entrance was black. Although there was no one apparent

overriding characteristic that allowed the Trap n' TossTM to capture more house flies than the

Terminator Pro, these studies document the variety of issues impacting house fly trapping.

Attractant Experiments

Two commonly used materials in commercial fly trap attractants are yeast and ammonium

carbonate. Pickens (1995) devised a categorizing system for the different odor attractants for

house flies. They are sugars and fermentation products, proteins, and animal excretions. Yeast

is in the sugar and fermentation products category, while ammonium carbonate is in both the

sugar and fermentation products as well as animal excretions category (Pickens 1995).

Cohen et al. (1991) used fly traps baited with yeast around a military camp and reported a

64% decrease in house fly density that accounted for a 42% decrease in clinic visits by the

soldiers for diarrhea and an 85% reduction in Shigellosis visits. Chapman et al. (1998) found

that yeast provided slightly increased house fly attraction over the control, while adding yeast to

(Z)-9-tricosene-impregnated targets produced a significant reduction in the number of male

house flies attracted. In my studies, I found yeast to be more attractive than the water control at

0, 2, and 4 days old by -30-35%. My studies do agree with both authors that yeast attracts more

house flies than a control, but my studies put yeast's attractiveness somewhere between the

attractancy observed by either Cohen et al. (1991) or Chapman et al. (1998).

Ammonium salts have been shown to be attractive to various filth flies most likely due to

the salts' release of ammonia. Ammonium carbonate is an ammonium salt and releases both

ammonium carbonate and carbon dioxide (Loeser et al. 2004). Mulla et al. (1977) lists

ammonium chloride and ammonium sulfate, as highly attractive to house flies. Wieting and









Hoskins (1939) found ammonia attractive and carbon dioxide not attractive to house flies.

Hobson (1935, 1936) found ammonium carbonate in conjunction with indole was attractive to

Lucilia sericata (Meigen). Mulla and Ridsdill-Smith (1986) found that when ammonium

carbonate was added to an attractant mixture, attractancy was not significantly increased for

Musca vetustissima. My studies demonstrated that ammonium carbonate was never more

attractive than the control, and when added to yeast (an attractant) did not increase attractancy.

The results that I obtained were never directly comparable with the aforementioned studies due

to the difference in attractant components used and/or species tested against. However, my

studies agree most with Mulla and Ridsdill-Smith's (1986) experiments withM. vetustissima, in

that ammonium carbonate does not represent an attractant chemical for the house fly when used

alone or in conjunction with other attractants.

Initially, there was reason to believe that a combination of yeast and ammonium carbonate

would be more attractive than the yeast or ammonium carbonate alone. Cosse and Baker (1996)

used Eletctoantennagram (EAG) to determine a number of chemicals that illicit a response in

house flies, but many required combination with other attractants to stimulate a behavioral

response i.e. attraction. Geden (2005) combined two commercial attractant mixtures and

reported synergistic effects. When yeast and ammonia carbonate were mixed, the combination

did not catch significantly more house flies than the yeast alone, but did catch more than

ammonium carbonate alone. There was certainly no synergistic effect. Because the assumed

attractive ammonium carbonate turned out to be not attractive, my results do not agree or

disagree with Cosse and Baker (1996) or Geden (2005).

Given the nature of the presumed attractant combination used (yeast and ammonium

carbonate components), it is logical to assume that the attractant combination would change over









time. Ammonium carbonate quickly released ammonia and carbon dioxide when added to water.

The yeast was a living organism and its life cycle was subject to availability of nutrition. In my

experiment, there was not an ample carbon supply for the yeast i.e. sugar, meaning the yeast

either arrested in GO phase, progressed slowly through the life cycle, or died (Dickinson and

Schwiezer 1999). How the yeast changed is unknown, but it was changing. Pickens et al. (1973)

stated that baits of fermented grains and yeasts took about three to four days to become

attractive. Geden (2005) observed commercial attractants to become less attractive by the fourth

day. The attractant combination in my study increased linearly from day 1 to 4 and agrees more

with Pickens et al. (1973), most likely due to the similarity of fermentation product attractants

that we each used. The components of the attractant mixtures used by Geden (2005) were not

reported; therefore it is difficult to discuss the similarities and differences. Also the experiments

of Geden (2005) were run outdoors at temperatures as high as 370 C, which may have affected

volatility or stability of the attractants, while my attractants were aged in less extreme

temperatures indoors (22.8 + 0.02 C) which may have kept the yeast viable.

Attractants increase the effectiveness of fly traps considerably. In fact, Kuzina (1940)

stated that odor was the most important mechanism used by house flies to locate food sources.

In my studies, yeast proved to be much better at attracting house flies than ammonium carbonate,

and ammonium carbonate never attracted significant numbers of house flies. The fact that

ammonium carbonate was not attractive, resulted in the combination of yeast and ammonium

carbonate being no better at attracting flies than the yeast alone. When using a live organism

such as yeast in order to attract house flies, as long as it is sustained, the attraction will increase

linearly for at least 4 days.









Conclusions

House flies are important pests that are best controlled by using IPM practices. This is

particularly true given the house fly's ability to develop insecticide resistance and its

considerable population potential (Hodge 1911, Scott et al. 2000, Learmount et al. 2002, Gao

and Scott 2006). Commercial fly traps have two potential niches in a house fly IPM program,

surveillance and control.

Field trials must still be run in order to obtain a better understanding on how well the tested

commercial fly traps perform in non-laboratory conditions. However my laboratory studies

demonstrated that fly traps can be viable devices for catching house flies. Although the Fly

Terminator Pro with an un-aged attractant mixture of yeast and ammonium carbonate only

captured -25% of the total house flies, the Trap n' TossTM with the same attractant mixture aged

4 days, caught -70% of the total house flies, and almost 80% of the female house flies. A

capture of -70% rivals the effectiveness of some biological control agents and insecticides.

Aside from comparing with other control measures, -70% capture exceeds Dadour and Cook's

(1992) requirement that in order to reduce a house fly population by 50-90%, 24-58% of the

present house flies should be captured. According to my laboratory studies, the proper design

(e.g. at least a 176.24cm2 entrance area, at most an 11.95 cm2 exit area, a ratio of at least 14.75, a

slope around 60, and a yellow entrance) and an aged attractant can capture high numbers of

house flies and should be considered for use as a control measure in an IPM program.

In addition to controlling fly populations, commercial fly traps may serve as monitoring

tools in a house fly IPM program. Burg and Axtell (1984) suggested that fly traps represented a

simple, practical way to monitor house fly populations, thus giving a better idea of when to use

additional control measures. Therefore, the commercial fly traps that did not catch the most

house flies in these studies could still be important tools in house fly IPM.









In conclusion, my studies have shown that commercial fly traps have potential as house fly

IPM tools. They are versatile devices to be used in IPM programs, and can be used either as

control devices or monitoring house fly populations. They are compatible with both insecticides

and biological control measures. Also, if there exists a situation in which insecticides or

biological controls cannot be used, they are safe, easy, and effective enough to be used as stand-

alone devices. According to my laboratory studies, and with the further exploration with field

studies, commercial fly traps represent a viable tool for managing house flies in a wide variety of

settings including agriculture, urban, and military









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BIOGRAPHICAL SKETCH

Ryan Merrill Welch was born on June 23, 1982, to Dwight and Amy Welch, in

Jacksonville, FL. There he attended Parkwood Heights Elementary School, Arlington Middle

School, and Stanton College Preparatory School. He graduated from Stanton College

Preparatory School with high school and international baccalaureate diplomas in 2000. He

attended the University of Florida from 2000 to 2004 and graduated cum laude earning a

Bachelor of Science in entomology and a Bachelor of Arts in political science. He attended the

University of Florida from 2004 to 2006 and received his Master of Science in entomology under

Dr. Phil Koehler's guidance.