Journal of Integrated Pest Management, (2021) 12(1): 39; 1–38

doi: 10.1093/jipm/pmaa021
Profile

House Fly (Diptera: Muscidae): Biology, Pest Status,

Current Management Prospects, and Research Needs
C. J. Geden,1,9, , D. Nayduch,2, J. G. Scott,3 E. R. Burgess IV,4 A. C. Gerry,5,
P. E. Kaufman,6, J. Thomson,7 V. Pickens, 7 and E. T. Machtinger8,
1
USDA, ARS, Center for Medical, Agricultural and Veterinary Entomology, 1600 SW 23rd Dr Gainesville, FL 32608, 2USDA, ARS, Cen-
ter for Grain & Animal Health Research, Arthropod-Borne Animal Diseases, 1515 College Ave, Manhattan, KS 66506, 3Department of
Entomology, 6134 Comstock, Cornell University, Ithaca, NY 14853, 4Department of Biological Sciences, Northern Illinois University,

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DeKalb, IL 60115-2865, 5Department of Entomology, University of California, 900 University Ave, Riverside, CA 92521, 6Department
of Entomology, 2475 TAMU, Texas A&M University, College Station, TX 77845, 7Department of Entomology, Kansas State University,
Manhattan, KS 66506-4801, 8Department of Entomology, Pennsylvania State University, 4 Chemical Ecology Laboratory University
Park, PA 16802, and 9Corresponding author, e-mail: chris.geden@ars.usda.gov

Subject Editor: Danesha Carley

Received 24 August 2020; Editorial decision 13 October 2020

Abstract
The house fly, Musca domestica L. (Diptera: Muscidae), is a global pest of humans and animals that carries scores
of pathogens and costs up to $1 billion per year in the United States alone. Information is reviewed on recognition,
distribution, biology, dispersal, and associations with microbes. Particular challenges of managing flies in different
animal systems are discussed for swine, poultry, dairy cattle, beef feedlot, and equine operations. Effective fly
management requires diligent monitoring and integration of cultural control, especially manure management,
with mechanical control, traps, conservation or augmentative biological control, and judicious use of insecticides.
House fly is notorious for developing insecticide resistance and its resistance status is summarized as of August
2020. Several critical research needs are identified. Monitoring systems and nuisance/action thresholds need
improvement. Faster-killing strains and better formulations are needed to integrate pathogens into Integrated Pest
management (IPM) programs. The use of parasitoids remains an inexact science with many questions remaining
about species selection and release rates. New attractants are needed for use in traps and attract-and-infect/kill
strategies. Screening of new active ingredients for toxicity should continue, including a rigorous assessment
of essential oils and other botanicals. Rising global temperatures may affect the balance of the fly with natural
enemies. An understanding of the fly microbiome may reveal unknown vulnerabilities, and much remains to be
learned about how flies acquire, retain, and transmit human and animal pathogens. System-specific research is
also needed to tailor fly IPM programs to individual animal systems, especially in organic and free-range animal
production.

Keywords: Musca domestica, Housefly, filth flies

House flies have been pests of humans and animals since an-
tiquity. Recent research by Gogarten et al. (2019) suggests that
the relationship between humans and flies may predate recorded
history for many millennia. These authors reported that muscid
and calliphorid flies are closely associated with social groups of
highly mobile nonhuman primates in the tropical forests of Ivory
Coast. The flies appear to move with the primates and are rarely
found outside of their immediate vicinity. Moreover, the flies carry
and presumably spread pathogens that cause disease in the associ-
ated animals. The authors conclude that ‘attraction of flies might
represent a previously underappreciated cost to forming social
groups’. The association between humans and flies was undoubt-
edly strengthened once humans began to form more permanent
Published by Oxford University Press on behalf of Entomological Society of America 2021. This work is written by (a) US Govern-
ment employee(s) and is in the public domain in the US.
settlements with domesticated animals and the concomitant ma-
nure accumulations. House flies were spread by humans as they
radiated across the planet and are found on every continent ex-
cept Antarctica. Their long synanthropy has allowed house flies
to adapt to a wide variety of environmental conditions and food
resources. This adaptability has resulted in the evolution of a for-
midable adversary that can avoid, adapt to, and evolve resistance
to our best efforts to bring it under control. The objective of this
article was to review the biology, pest status, and current manage-
ment prospects for this important pest and to suggest areas where
future research is needed.
1
2 Journal of Integrated Pest Management, 2021, Vol. 12, No. 1

Recognition and Distribution

Adult house flies have reddish eyes, sponging mouthparts, are
3–8 mm in length and can be recognized by the presence of four
dark stripes on the dorsum of the thorax and the pronounced
upward bend in the fourth longitudinal wing vein (vein M1+2)
(Figs. 1 and 2). The basal portion of the abdomen is usually
yellowish, especially along the sides, with males typically showing
greater lateral yellowing than females. A dark longitudinal band
runs along the median dorsal region of the anterior abdominal seg-
ments. The eyes of females are much more widely separated than
those of males (Fig. 3). The key provided in Crosskey and Lane
(1993) is useful for distinguishing Musca domestica from related
muscoid flies. Eggs are deposited in batches in moist substrates,
are creamy white, and approximately 1 mm long (Fig. 4). Mature
house fly larvae can be recognized by the ‘D’-shaped peritreme

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surrounding sinusoidally shaped spiracular slits (Fig. 5). Pupae
range in color from light red to dark brown, are 4–7 mm long, and
have hardened spiracles with the same characteristics as the larvae
(Fig. 6). House flies are found most commonly and abundantly
at animal production facilities, but also occur in urban settings
Figure 2. Side view of female house fly head showing sponging mouthparts.
where the larvae develop in a wide range of decaying vegetable
Photo by Matt Aubuchon.
materials, feces, and household garbage. The common denomin-
ator among suitable developmental substrates is an abundant, vi-
Feeding
able microbial community.
Adult house flies have sponging mouthparts and therefore must con-
sume either liquid food or regurgitate crop contents onto solid foods
General Biology to soften them prior to ingestion. ‘Bubbling’ is a common phenom-
enon, in which flies exude a droplet of regurgitate. Bubbling is thought
House fly biology, life history, and management have been re-
to eliminate excess water (Hendrichs et al. 1992, Stoffolano 2019),
viewed often and extensively. Early researchers developed a sound
thus concentrating the nutrients in the ingested food and reducing the
understanding of the fly’s biology over 100 yr ago (Howard 1911,
weight that flies must carry during flight. Bubbling may also play a role
Graham-Smith 1913, Hewitt 1914a), and the volume by West
in thermoregulation (Gomes et al. 2018). The droplets are sometimes
(1951) remains an essential work for its excellent anatomical illus-
deposited in the environment, leading to accumulations of both regur-
trations and overview of critical life history events. Greenberg (1971,
gitation and fecal spots on surfaces where flies rest (Fig. 7). The relative
1973) assembled the known literature on the association of flies and
amount of regurgitation spots varies with the quality of the food in-
disease organisms into a 2-volume set. Since then, there have been
gested. Flies that have fed on nutrient-dense foods such as blood, liquid
several reviews with different emphases on house fly biology, physi-
milk, and high-concentration sugar solutions produce fewer regurgi-
ology, management, or ecological associations (Keiding 1986, Axtell
tation spots than flies fed on items that are of lower nutrient density
and Arends 1990, Axtell 1999, Hogsette and Farkas 2000, Malik
(CJG, unpublished data). Deposition of regurgitation and fecal spots
et al. 2007). Published proceedings of two national workshops on
play a role in the transmission of pathogens and can be used in moni-
research and extension needs for veterinary entomology included
toring programs as an index of fly abundance.
house fly information in chapters addressing fly concerns in different
The crop, a ventral diverticulated structure of the foregut, pro-
animal production systems (Anonymous 1976, Geden and Hogsette
vides the fly with a highly expandable storage organ that allows it
2001). The following is intended as an overview of basic house fly
to consume large quantities of high-quality liquid food when such
life history.
foods are encountered (reviewed in Stoffolano and Haselton 2013,
Stoffolano 2019). Crop contents are then either passed to the midgut
for digestion or passed forward for release from the proboscis as re-
gurgitate. Food sometimes bypasses the crop and goes directly into
the midgut, especially when ingested food is low in moisture content.
As food enters the midgut it is surrounded by the peritrophic matrix
(PM). The PM is synthesized by midgut cells and is composed of a
matrix of chitin, glycoproteins, peritrophins, and other components
that provide a barrier that prevents ingested pathogens from infecting
the fly by denying them contact with the microvillar membrane of the
midgut epithelium. Nutrients pass through the PM and are absorbed
in the midgut, after which any unprocessed food enters the hindgut
for water absorption and eventual deposition as feces. Passage of food
through the entire digestive system can be completed in less than 6 h.

Reproduction
Figure 1. Female house fly. Arrows indicate distinct upward bend in the Female flies generally mate once, usually within 36 h of eclosion.
fourth longitudinal wing vein. Photo by Matt Aubuchon. A putative sex pheromone, (Z)-9-tricosene, was identified as a
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1 3

Figure 3. House flies (5–8 mm) have a dull gray thorax with 4 thoracic stripes. The female (right) has widely separated eyes and a lightly golden checkered

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abdomen. The male’s eyes are positioned close to each other and has a darkly checkered abdomen. Photo by Matt Bertone.

Figure 5. The creamy white third instar (5–8 mm) of the house fly with the
blackened, internal mouth-hooks in the front and the distinctive D-shaped
Figure 4. House fly eggs are small (~1 mm), white, and deposited on or just spiracles with sinusoidal slits at the rear. Graphic courtesy of Fallon Fowler
under the surface of moist substrates. Photo by Steve Denning. using photos by Matt Bertone and Steve Denning.

cuticular hydrocarbon constituent of female house flies (Carlson

et al. 1971). This material is attractive to flies and is often incorp-
orated into baits but is not present in all populations of flies col-
lected in the field (Darbro et al. 2005, Butler et al. 2009). Amounts
of (Z)-9-tricosene are higher in laboratory-reared flies than in wild
flies (Noorman and Den Otter 2001, Darbro et al. 2005) and in-
crease with fly age (Butler et al. 2013). Flies under laboratory con-
ditions and given highly nutritious food are ready to deposit their
first egg batch within 3–5 d after emergence and can lay 100–150
eggs per female per gonotrophic cycle. Because flies under such Figure 6. The reddish-brown puparium (4-7mm) of the house fly with
hardened spiracles at the rear. Photo by Matt Bertone.
conditions can produce up to six batches of eggs, an individual fe-
male has the potential to lay 900 eggs in her lifetime (West 1951).
Fletcher et al. (1990) observed that flies laid a maximum 709 eggs
per female at 30°C under laboratory conditions with adequate food. Longevity
Protein is required for egg maturation, but flies can develop eggs Adult female flies held under optimal laboratory conditions can sur-
even when allowed brief, intermittent bouts of feeding on a protein vive up to 6 wk (Fletcher et al. 1990, Haselton et al. 2004). Field esti-
source (Adams and Gerst 1991). Under field conditions, where fly mates of fly longevity, based on mark-release-recapture methods, are
longevity is shorter and food resources often limiting, rates of egg de- much shorter than estimates from laboratory studies. The lifespans of
position are undoubtedly much lower. Lysyk (1991a) observed that wild flies have been estimated at 1–6 d and 3–19 d for flies on poultry
flies given poultry feed and manure deposited only about 40 eggs per (Lysyk and Axtell 1986c, Lysyk 1991a) and dairy farms (Krafsur et al.
female over a 9- to 12-day lifespan, and Krafsur (1985) estimated 1985, Kristiansen and Skovmand 1985, Butler et al. 2013), respect-
that females on an Iowa dairy farm produced an average of 94 eggs ively, and 2–7 d for flies collected from a refuse dump (Imai 1984).
over an approximate lifespan of 5.4 d. Oviposition occurs in groups These results suggest that some production systems favor fly lon-
and is stimulated by semiochemicals from the ovaries of gravid fe- gevity over others. The short lifespans of flies reported from land-
males (Jiang et al. 2002) and can be further modulated by bacterial fills and poultry farms may reflect poor food resources at such sites,
symbionts deposited along with the eggs (Lam et al. 2007). whereas dairies offer a wider variety of high-nutrient foods such
4
Figure 7. Fly fecal and regurgitation spots accumulate where flies rest in the environment. Photos by Erika Machtinger.
as milk, occasional blood from animal wounds, and pelleted feeds
given to young animals.
Immature Development
Development time is determined by temperature and the quality of
the larval substrate. Eggs generally hatch 6–12 h after deposition.
Larsen and Thomsen (1940) found that development from egg to
adult was completed in less than 7 d at 33°C. Lysyk and Axtell
(1987) reported that the average time from oviposition to pupation
ranged from 26.8 d at 16°C to 5.2 d at 35°C, and that the average
time to adult emergence ranged from 43.1 to 8.8 d. Larvae move
to dry areas to pupate and the duration of the pupal stage is 3–6 d
under summer conditions. It is worth noting that development time
experiments based on laboratory colonies may result in substantial
underestimates. Early generations of new colonies from field collec-
tions have longer larval development times and higher pupal weights,
and are much more variable in development times than colonies that
have adapted to laboratory conditions (CJG, unpublished data).
The quality of the larval substrate also modulates development
time. Larvae that are reared in suboptimal substrates often complete
development but require longer larval stadia. Fly larvae reared in
horse manure can take 50% longer to develop than larvae reared
in poultry manure (Khan et al. 2012). Hogsette (1996) found that
house fly larval development at 26.7°C required 22 d when they were
reared in moist sand containing small amounts of dairy cattle ma-
nure. House fly larvae also can develop in a wide range of decaying
plant substrates, including crop residues and culls after harvesting
such as corn, carrots, onions, and snow peas (Cook et al. 2011).
Regardless of the substrate, fly larvae must consume live micro-
organisms to complete their development (discussed below under
fly-microbe associations).
Dispersal
On agricultural installations where house fly breeding typically
occurs, most flies remain on or near the facility (Lysyk and Axtell
1986b). They are, however, capable of moving longer distances and
can create problems when they disperse from the production site to
residential areas, schools, and businesses (Thomas and Skoda 1993,
Lole 2005, Winpisinger et al. 2005). Dispersal distances of >12 km
have been reported (Bishopp and Laake 1921, Quarterman et al.
1954, Greenberg 1973), and Yates et al. (1952) collected a fly 32
km from the release point. Such reports of long-distance movements
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should be viewed with caution, however, since the vast majority of
flies remain close to where they emerge. Dispersal distances vary de-
pending on food availability and/or the number of development sites
encountered by flies as they move through their environment (Schoof
et al. 1952, Schoof and Siverly 1954a,b, Schoof 1959). Habitat
structure can provide corridors for longer distance travel (Fried et al.
2005). In summary, most house fly movement from agricultural fa-
cilities is likely a series of short disjointed circuitous flights culmin-
ating in the aggregation of house flies at some more distant attractive
site (Schoof and Siverly 1954a,b) rather than a directional dispersal
to discover and colonize new habitats. It is notoriously difficult to
determine the source populations of flies when problems and com-
plaints arise on nonfarm sites, often resulting in disputes between
animal operations, neighbors, and local health officials.
Genetics
The house fly has five pairs of autosomes plus an X and Y chromo-
some. Sex is determined by the Mdmd gene (Sharma et al. 2017),
which can be found on any of the chromosomes and can be pre-
sent in more than one copy (Hamm et al. 2015, Son et al. 2019).
Y is the presumed ancestral chromosome for Mdmd. The house fly
genome was sequenced in 2014 (Scott et al. 2014) and revealed an
abundance of recognition and effector components of the immune
system, consistent with its close association to pathogens. Initial
studies found a lack of crossing-over in male house flies (McDonald
1971), consistent with what is observed in most Diptera (Gethmann
1988). Subsequently, it was found that crossover frequencies in
males vary, depending on the genes examined and the populations
used. Reported values range between 0–0.53% (Hamm et al. 2005,
Hamm and Scott 2008), 0.03–0.11% (Sullivan 1961), 9.3–31%
(Lester et al. 1979), and 7–28% (Feldmeyer et al. 2010). Greater
male recombination rates tend to be associated with males that have
Mdmd on an autosome.
Fly–Microbe Interactions
Larval Associations
Larvae feed on live microorganisms within the substrate. Gerberich
(1948) and Greenberg (1954) first noted that house fly larvae must
ingest living microorganisms to complete development and suggested
that this was due to B vitamins or other nutritional factors provided
by gut-inhabiting microbes. When grain-based fly larval diets were
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1 5

first developed, it was recommended that they be inoculated with
a cocktail of Escherichia coli, Sarcina spp., and Lactobacillus spp.
to promote larval development (Spiller 1964). Some early studies
suggested that house fly larvae could be grown under axenic condi-
tions (Brookes and Frankel 1958, Monroe 1962), but the methods
may not have been sufficiently sterile to rule out colonization by
microbes. House fly larvae grown on blood agar without bac-
teria do not develop past the first instar, but can complete devel-
opment on monoxenic plates with E. coli, Klebsiella pneumoniae,
Staphylococcus spp., and Streptococcus bovis (Schmidtmann and
Martin 1992). Similarly, Watson et al. (1993) successfully grew
house fly larvae on sterile egg yolk and blood agar media inocu-
lated with E. coli, whereas almost no adult flies were produced on
sterile media. Bacteria isolated from the gut of fly larvae in soiled
turkey litter, including the Gram-positive facultative anaerobic spe-
When flies ingest bacteria, the location and persistence of the
bacteria within the digestive tract impacts transmission potential.
Ingested bacteria harbored in the crop and midgut are either digested
and destroyed or survive to be excreted and, possibly, transmitted
(Nayduch and Burrus 2017, Stoffolano 2019). Bacterial persistence,
propagation, excretion, and transmission can vary by pathogen
type (Nayduch et al. 2002, Joyner et al. 2013, Nayduch et al. 2013,
Fleming et al. 2014). Even fly sex can impact the acquisition, per-
sistence, proliferation, and excretion of pathogens (Thomson et al.
2017, Nayduch et al. 2018).
Transmission of pathogens by flies to human food items is of
growing concern in light of several high-profile recalls of contam-
inated leafy greens (Talley et al. 2009). Transmission of disease-
causing E. coli from house flies to spinach leaves was shown by
Wasala et al. (2013). Viable cells of E. coli O157:H7 were detected

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cies Staphylococcus lentus, Streptococcus sanguinis, Lactococcus
garviae, Yersinia pseudotuberculosis, and Bacillus coagulans all
supported house fly development in monoxenic trypticase soy egg
yolk agar; larval development was poor when the bacteria were ones
recovered from larvae found in fermented corn silage (Zurek et al.
2000). The authors concluded that ‘house fly larvae likely benefit
from complex metabolic interactions within a diverse bacterial com-
munity in a natural environment leading to a rapid degradation of
organic material as well as a great build-up of bacterial mass’. The
house fly larval microbial community is complex and variable, and
horizontal transmission of microbes among larvae in feeding sites
ensures a continuous local supply of organisms (Zhao et al. 2017).
Acquisition, Retention, and Transmission of
Microbes by Adult Flies
Adult house flies routinely acquire microbes such as protists, viruses
and bacteria during their persistent associations with microbe-rich
environments. Over 200 different species of microbes have been iso-
lated from wild-caught house flies (Nayduch and Burrus 2017), and
a single house fly can carry up to 100 different pathogenic microbes
(Greenberg 1973). Flies acquire microbes on their surfaces via contact
with or by directly feeding upon refuse, animal waste, wounds and
exudate (West 1951, Nayduch and Burrus 2017). Pathogens on flies’
surfaces are transferred to animals by physical contact or by being
dislodged by fly grooming (Yap et al. 2008, Jacques et al. 2017). Some
bacteria such as E. coli O157:H7 survive and proliferate for several
days on mouthparts before transmission (Kobayashi et al. 1999).
8 d after contaminated flies contacted the spinach. Because E. coli
cells cannot survive 8 d, the flies must have transmitted viable cells
(Wasala et al. 2013). Transmission by house flies has been shown
for a number of bacterial pathogens under a variety of test con-
ditions, including contamination of the environment and human
food items (Tables 1 and 2).
Flies potentially facilitate the evolution and dispersal of new
pathogenic and resistant microbial strains (Petridis et al. 2006,
Onwugamba et al. 2018). For example, lateral gene transfer be-
tween bacteria may occur in the fly gut (midgut + crop), including
the horizontal movement of resistance alleles among pathogens
and nonpathogens (Akhtar et al. 2009, Zurek and Ghosh 2014,
Onwugamba et al. 2018, Poudel et al. 2020). The horizontal transfer
of a plasmid harboring cephalosporin and tetracycline resistances
between donor and recipient bacteria in the house fly gut has been
demonstrated (Fukuda et al. 2016). The plasmid was not only trans-
ferred between donor and recipient E. coli strains, but also to other
bacterial taxa that were present as ‘normal flora’ (Achromobacter
spp. and Pseudomonas fluorescens). The horizontal transfer of plas-
mids carrying tetracycline resistance among enterococci has also
been demonstrated within the house fly gut (Akhtar et al. 2009).
Furthermore, flies may facilitate the introduction and dispersal of
genes affecting virulence (Zurek and Ghosh 2014). Because adult
flies are highly mobile, they substantially contribute to the dissemin-
ation of these novel pathogens across habitats and ecological niches
(Zurek and Ghosh 2014, Onwugamba et al. 2018, Poudel et al.
2019, Sobur et al. 2019).

Table 1. Studies demonstrating transmission of pathogenic bacteria by house flies

Pathogen Extent of dissemination References

Aeromonas caviae Contaminate environment Nayduch et al. 2002

Aeromonas hydrophila Viable in excreta McGaughey and Nayduch 2009
Campylobacter jejuni Viable in excreta Gill et al. 2017
Contaminate environment Shane et al. 1985
Corynebacterium pseudotuberculosis Viable in excreta Braverman et al. 1999
Enterococcus faecalis Contaminate environment Doud and Zurek 2012
Escherichia coli O157:H7 Viable in excreta Sasaki et al. 2000, Fleming et al. 2014
Contaminate environment Wasala et al. 2013
Pseudomonas aeruginosa Viable in excreta Joyner et al. 2013
Salmonella Typhimurium Viable in excreta Chifanzwa and Nayduch 2018
Salmonella Schottmullerris Viable in excreta Hawley et al. 1951
Salmonella Enteritidis Transmit to hens Holt et al. 2007
Shigella dysenteriae Viable in excreta Hawley et al. 1951
Staphylococcus aureus Viable in excreta Nayduch et al. 2013
Yersinia pseudotuberculosis Contaminate environment Zurek et al. 2001
6
Dissemination of Pathogens by Adult Flies
Adult flies bridge unsanitary and sanitary environments,
disseminating bacteria from their source to animal facilities, food,
water, and nearby humans (Nayduch and Burrus 2017) (Fig. 8).
Fly populations flourish in livestock facilities and the threat to
human and livestock health is intensified when poor sanitation
and insufficient manure management allows flies to have unre-
stricted access to pathogen sources such as waste and excrement.
House fly dispersal between farms and nearby residential and
urban centers facilitates bacterial transmission to humans and
therefore poses a public health risk. As discussed previously, flies
can disperse several kilometers from their larval habitats (e.g.,
farms). This increases the potential to spread manure-acquired
bacteria, including antimicrobial resistant (AMR) strains, to sur-
rounding locales (Alam and Zurek 2004, Winpisinger et al. 2005,
Burrus 2010, Baldacchino et al. 2018, Neupane et al. 2020).
Chakrabarti et al. (2010) estimated that flies could facilitate
movement of antibiotic-resistant bacteria as far as 100 km from
a cattle feedlot. Flies may spread AMR E. coli among animal pro-
duction sites that are geographically separated (Usui et al. 2015),
which may contribute to multiple AMR E. coli strains occurring
within single facilities (Sola-Gines et al. 2015). Other specific ex-
amples of fly transmission include dispersal of methicillin-resistant
Staphylococcus aureus to urban communities (Schaumburg et al.
2016), transmission of AMR serotypes of various Salmonella spp.
within and among swine farms (Wang et al. 2011), transmission
of Corynebacterium pseudotuberculosis, a causative agent of bo-
vine mastitis (Yeruham et al. 1996), and possibly dissemination of
Table 2. Food-borne pathogens transmitted to or from food by
house flies
Pathogen Food type References
Aeromonas caviae Various Nayduch et al. 2002
enterococci Cooked hamburger Macovei et al. 2008
patty
Escherichia coli O157:H7 Spinach Wasala et al. 2013
Lettuce Pace et al. 2017
Antimicrobial-resistant Various Fukuda et al. 2019
E. coli
Salmonella enteric Mexican drink Greenberg 1964
Lettuce Pace et al. 2017
Canteloupe Thomson et al. 2020
Only includes studies that have specifically described food-borne pathogens
and food sources and demonstrated house fly impact on food safety.
Figure 8. Flies aid in transmission of animal pathogens when they visit animal manure and food resources. Photos by Chris Geden.
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
Campylobacter (Hald et al. 2007) or Salmonella spp. (Olsen and
Hammack 2000) at poultry facilities.
House Fly Impacts in Human and
Animal Systems
Economic Losses
It is difficult to determine the present economic scope of the house
fly problem. After adjusting for inflation, a 2001 estimate for the
annual cost of insecticides for fly control in the poultry industry is
about $30 million today (Geden et al. 2001). Estimates for the dairy
and swine industries, after adjusting for inflation, are $135 million
and $35 million, respectively (USDA 1978, Campbell 1993). Taken
together, the cost of insecticide use for house flies in these three com-
modities can therefore be estimated at about $200 million. A 1976
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estimate for the total cost of house fly damage and control is ap-
proximately $450 million today (Anonymous 1976).
Even after adjusting for inflation, these figures are undoubtedly
low. One reason for this is the high cost of legal settlements against
producers in recent decades. Changing demographics have resulted
in growing friction between once-isolated agriculture facilities and
residential developments. House flies were one of the complaints in
the recent $50,000,000 million settlement against Smithfield Foods
for nuisance complaints regarding hog farms (Brown 2018). Societal
and public health pressures make it necessary to keep fly population
levels lower than were acceptable when flies were seen as being a
natural outcome of living in rural settings. Moreover, the relatively
higher cost of modern insecticides has made fly control more ex-
pensive than in past years. Until more solid economic data become
available, we estimate that house flies account for $500 million to
$1 billion per year in insecticide costs and total economic losses,
respectively.
Human Health Concerns
Due to their persistent associations with decomposing substrates,
flies carry a rich and diverse bacterial community. House flies are
known to carry an ever-growing list of human pathogens that can be
transmitted directly to people or indirectly via contamination of food
(Sundin 1996, Graczyk et al. 2001, Boulesteix et al. 2005, Rahuma
et al. 2005, Macovei and Zurek 2006, Macovei et al. 2008, Graham
et al. 2009, Nayduch and Burrus 2017, Khamesipour et al. 2018, Xu
et al. 2018). Infected flies often originate from concentrated livestock
operations, which generate large numbers of house flies that have
access to potential zoonotic and AMR pathogens shed in animal ma-
nure. Many different pathogenic bacteria have been isolated from
livestock-associated flies, including AMR strains and both human
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
and livestock pathogens (Table 3). Other pathogens such as viruses,
protists, and helminth eggs are also associated with wild house flies,
albeit only transiently and in less abundance; they are reviewed else-
where (e.g., Graczyk et al. 2001, Nayduch and Burrus 2017).
Flies pose a risk to humans living near sources of human patho-
genic microbes including livestock and poultry operations, landfills,
and wastewater management facilities (Iwasa et al. 1999, Zurek
and Ghosh 2014, Schaumberg et al. 2016). Several studies have
suggested flies as a source of human pathogenic bacteria in animal
operations, including E. coli O157:H7 (Rahn et al. 1997, Hancock
et al. 1998, Iwasa et al. 1999, Alam and Zurek 2004, Szalanski et al.
2004), Camplylobacter spp. (Szalanski et al. 2004, Ekdahl et al.
2005), and Salmonella spp. (Mian et al. 2002). A recent survey of
house flies from dairies and feedlots in Georgia identified Salmonella
spp., including strains with AMR, in 11% (185/1650) of flies and
pathogen incidence varied widely across sites (0–78%) (Xu et al.
2018). Single-drug AMR also was reported in some of these isolates,
and 28% of the Salmonella isolates were multidrug resistant. Flies
carrying enteropathogenic E. coli which originated from a nearby
animal farm were posited as the source of a colitis outbreak at a
children’s school in Japan (Moriya et al. 1999). House flies collected
from wastewater management facilities carried antibiotic resistant
Enterococcus faecalis and a few flies were found to carry these re-
sistant bacteria in a recreational vehicle park, a fast food restaurant,
and an apartment complex close to some of the treatment facilities
(Doud et al. 2014).
Flies also pose a risk when they have access to human garbage and
feces in situations where basic sanitation practices are poor. House
flies were connected to outbreaks of typhoid fever from Salmonella
typhi during the Spanish-American war in military camps (Cirillo
2006) and a dysentery outbreak in a U.S. army camp (Kuhns and
Anderson 1944). The proximity of food and water sources to feces
in addition to the presence of higher than average fly densities con-
tributed to a 15% increase in the risk of diarrhea in humans in parts
of India (Collinet-Adler et al. 2015). In Bangladesh, flies were im-
plicated in contaminating food with enteropathogenic E. coli that
originated from nearby human feces (Doza et al. 2018). The bazaar
fly, Musca sorbens Wiedemann, a close relative of house fly with a
strong affinity for human eye secretions, is an important vector of
the causative agent of trachoma (Chlamydia trachomatis) in many
developing countries (Emerson et al. 2000); M. domestica appears to
be a competent vector as well (Stoffolano, personal communication).
Although there is no current direct evidence of house flies being able
to transmit the Ebola or SARS-CoV-2 viruses, fly control is recom-
mended as a sanitation precaution against mechanical transmission
(Haddow et al. 2017, Dehghani and Kassiri 2020).
One way to demonstrate the role of house flies in transmit-
ting pathogens is by showing that lack of fly control is linked to
diarrheal disease prevalence. Shortly after the discovery of DDT,
a study in Texas found that the use of this insecticide for fly con-
trol reduced the prevalence of diarrheal diseases in children under
the age of 5 (Watt and Lindsay 1948). The prevalence of Shigella
infections was significantly lower only during the time when an ef-
fective fly control program was carried out in select towns of rural
Georgia (Lindsay et al. 1953). McCabe and Haines (1957) showed
that over the 18 yr after reconstructing outhouses in Boston to re-
duce house fly breeding in human feces, the frequency of Shigella
infections in children decreased and the diarrheal disease rate for
Boston was cut in half. In Bangladesh, a surge of house fly popu-
lations in the spring was correlated with the subsequent increase
in Shigella diarrhea among children two months later (Farag et al.
2013). Similarly, the incidence of diarrheal disease was lower in
7
the children from towns in Pakistan where insecticides were used
to control flies (Chavasse et al. 1999). Fly control at an Israeli
military base resulted in an 85% drop in clinic visits for shigellosis
(Cohen et al. 1991).
Animal Health Concerns
Filth flies, including house flies, are key players in food security (e.g.,
animal health) and both pre- and postharvest food safety across a
variety of livestock commodities (Mian et al. 2002, Dhillon et al.
2004, Winpisinger et al. 2005, Holt et al. 2007, Hald et al. 2004).
The list of pathogenic microbes found in association with flies, es-
pecially those associated with livestock, is vast and continues to
grow (Keiding 1986, Scott et al. 2014, Nayduch and Burrus 2017,
Khamesipour et al. 2018). House flies are mechanical vectors of
scores of pathogens responsible for animal diseases. Many of the
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bacterial species listed in Table 3 are animal pathogens.
Livestock diseases can have differing impacts depending on typ-
ology and severity. Diseases impacting the food supply can have pro-
found effects on the entire livestock industry as outbreaks of some
diseases can lead to dramatic supply reductions, partial, or full stop-
pages of varying duration with trading partners, as well as a hesita-
tion to consume meat products from regions that are found to have
livestock disease outbreaks. A less severe scenario includes a spread
of disease throughout the herd, causing diminished productivity or
an increase in mortality and morbidity. Commodity specific patho-
gens are discussed below.
Challenges for Different Animal Systems
Animal production systems are continuing a trend of greater in-
tensification to maximize production while minimizing costs that
started in the previous century (Machtinger et al. 2020). One
of the hallmarks of this intensification is that increased animal
density results in greater production of animal waste and a greater
need for (and often storage of) feed per unit of area (Gerry 2018,
Clay et al. 2020)). The abundance of suitable fly larval develop-
ment habitat, particularly animal manure, is the most important
determinant of house fly presence and populations sizes in such
systems. Manure is typically collected and stored at the facility,
providing an uninterrupted input of fly-development substrates
into the system. The mixture of manure with animal feed and bed-
ding is another feature of intensive confined animal production,
and these mixtures provide ideal fly development sites. Control of
house flies under these conditions is difficult as it requires substan-
tial effort to manage manure and other developmental substrates.
Swine, poultry, dairy cattle, beef feedlots, and equine systems have
many elements in common, but each poses particular challenges.
Some of these are outlined below, as well as examples of disease
concerns in each system where house flies play a documented or
suspected role.
Swine
Swine production has moved toward fewer farm numbers and in-
creased concentrations of animals over the past few decades (USDA
2017) (Fig. 9). These large numbers of swine produce waste and
decaying organic matter that may serve as development sites for
house flies. While understanding production and legal risks associ-
ated with house flies has increased over the past decades, informa-
tion on the muscoid fauna in swine facilities in the United States
is limited. Burns and Nipper (1960) reported that house flies were
the most important insect problem associated with pig parlors in
8 Journal of Integrated Pest Management, 2021, Vol. 12, No. 1

Table 3. Animal and human pathogens found in flies collected at different animal systems

Bacteria Animal system References

Dairy Beef Cattle Swine Poultry Equine Other

cattle cattle (unspecified)

Acinetobacter spp. ✓ Nazni et al. 2005

Acinetobacter ✓ Nazni et al. 2005
baumannii
Aeromonas spp.a ✓ ✓ ✓ ✓ Nayduch et al. 2001; Ommi et al. 2015a
Aeromonas caviae ✓ ✓ Nayduch et al. 2001
Bacillus spp. ✓ ✓ Nazni et al. 2005; Bahrndorff et al. 2017
Campylobacter spp.a ✓ ✓ ✓ Szalanski et al. 2004; Brazil et al. 2007; Choo
et al. 2011; Royden et al. 2016; Ommi et al.
2016a
Campylobacter coli ✓ ✓ Rosef and Kapperud 1983; Hald et al. 2008

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Campylobacter fetus ✓ ✓ Rosef and Kapperud 1983
subsp. jejuni
Campylobacter jejuni ✓ ✓ Hald et al. 2008; Förster et al. 2009a,b
Citrobacter spp.a ✓ ✓ ✓ ✓ Moissant et al. 2004, Neupane et al. 2020a
Clostridium spp. ✓ Bahrndorff et al. 2017
C. perfringens ✓ Dhillon et al. 2004
Coccobacillus spp. ✓ Nazni et al. 2005
Corynebacterium spp. ✓ Hernandez 2012
Corynebacterium ✓ Braverman et al. 1999
pseudotuberculosis
Edwardsiella spp. ✓ Shukla et al. 2013
Enterobacter spp. ✓ ✓ ✓ ✓ Moissant et al. 2004; Nazni et al. 2005;
Neupane et al. 2020a
Enterobacter ✓ Buma et al. 1999a
sakazakiia
Enterococcus spp. ✓ Graham et al. 2009a
Enterococcus ✓ Ahmad et al. 2011a
casseliflavusa
Enterococcus faecalisa ✓ ✓ Ahmad et al. 2011a
Enterococcus faeciuma ✓ ✓ Ahmad et al. 2011a
Enterococcus hirae ✓ Ahmad et al. 2011
Escherichia spp. ✓ ✓ ✓ Nazni et al. 2005; Nmorsi et al. 2007; Förster
et al. 2009a,b
Escherichia colia ✓ ✓ ✓ ✓ ✓ ✓ ✓ Echeverria et al. 1983; Oo et al. 1989; Iwasa
et al. 1999; Agui 2001; Alam and Zurek
2004; Moissant et al. 2004; Szalanski et al.
2004; Brazil et al. 2007; Nmorsi et al. 2007;
Literak et al. 2009a; Usui et al. 2013a; Usui
et al. 2015a; Blaak et al. 2014a; Solà-Ginés
et al. 2015a; Schaumberg 2016a; Bahrndorff
et al. 2017; Puri-Guri et al. 2017; Neupane
et al. 2020a
E. coli 0157:H7 ✓ ✓ ✓ ✓ Buma et al. 1999a; Iwasa et al. 1999; Szalanski
(EHEC)a et al. 2004; Burrus 2010; Förster et al.
2009a,b; Burrus et al. 2017
Histophilus somni ✓ Neupane et al. 2019
Klebsiella spp.a ✓ ✓ Nazni et al. 2005; Nmorsi et al. 2007; Neupane
et al. 2020a
Klebsiella ✓ ✓ ✓ Nmorsi et al. 2007; Ranjbar et al. 2016a;
pneumoniaea Neupane et al. 2020a
Lactobacillus spp. ✓ Hernandez 2012
Listeria spp. ✓ ✓ ✓ ✓ Hernandez 2012
Mannheimia ✓ Neupane et al. 2019
haemolytica
Micrococcus spp. ✓ Nazni et al. 2005
Pasteurella multocida ✓ Neupane et al. 2019
Proteus spp.a ✓ ✓ Nazni et al. 2005; Nmorsi et al. 2007; Neupane
et al. 2020a
Proteus mirabilis ✓ Nmorsi et al. 2007
Providencia spp.a ✓ Shukla et al. 2013; Neupane et al. 2020a
Pseudomonas spp.a ✓ ✓ Hemmatinezhad et al. 2015a
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1 9

Table 3. Continued

Bacteria Animal system References

Dairy Beef cattle Cattle Swine Poultry Equine Other

cattle (unspecified)

Pseudomonas ✓ Buma et al. 1999a

fluorescensa
Salmonella spp.a
Salmonella Heidelberg
Salmonella infantis
Salmonella typhi
Salmonella
✓ ✓ ✓ ✓ ✓ Oo et al. 1989; Olsen and Hammack 2000;
Nmorsi et al. 2007; Choo et al. 2011; Wang
et al. 2011a; Xu et al. 2018a
✓ Olsen and Hammack 2000
✓ Olsen and Hammack 2000
✓ Nmorsi et al. 2007
✓ Nmorsi et al. 2007

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typhimurium
Serratia spp.a ✓ Nmorsi et al. 2007; Neupane et al. 2020a
Serratia marcescensa ✓ Buma et al. 1999a; Neupane et al. 2020a
Shigella spp. ✓ ✓ ✓ ✓ Echeverria et al. 1983; Oo et al. 1989; Shukla
et al. 2013
Staphylococcus spp.a ✓ ✓ ✓ ✓ ✓ Nazni et al. 2005; Nmorsi et al. 2007; Graham
et al. 2009a; Hernandez 2012; Bahrndorff
et al. 2017
Staphylococcus ✓ ✓ ✓ Nmorsi et al. 2007; Schaumberg et al. 2016a
aureusa
Stenotrophomonas ✓ ✓ Fukuda et al. 2017a
maltophilaa
Streptococcus spp. ✓ ✓ Nazni et al. 2005; Nmorsi et al. 2007
Streptococcus faecalis ✓ Nmorsi et al. 2007
Streptococcus ✓ Nmorsi et al. 2007
pyogenes
Vibrio spp. ✓ ✓ ✓ ✓ ✓ Echeverria et al. 1983; Hernandez 2012
Vibrio cholera ✓ ✓ ✓ Oo et al. 1989

a
Antimicrobial resistance found in bacterial isolates from house flies.

Figure 9. Swine in the U.S. are usually held in high-density enclosed facilities.
Manure is collected into liquid storage systems of various types, but barn
design sometimes leaves corners and other places that are difficult to clean.
Source: USDA NCRS, photo by Bob Nichols.
Louisiana, whereas house flies accounted for only 11.8% of filth flies
in confined hog facilities in Texas (Robertson and Sanders 1979).
Porcine reproductive and respiratory syndrome (PRRS) virus,
one of the most economically significant pathogens in the swine in-
dustry (Holck and Polson 2003), can be transmitted by house flies
(Otake et al. 2003). PRRS is estimated to cost the U.S. swine in-
dustry approximately $560 million per year (Neumann et al. 2005).
House flies transmit PRRS among animals within a facility and can
move PRRS from one facility to another (Schurrer et al. 2004, Pitkin
et al. 2008). House flies may also play a role in between-farm move-
ment of porcine epidemic diarrhea virus (PEDV) (Masiuk et al. 2018,
Allison et al. 2019, 2020). PEDV first emerged in the United States in
2013, and by May 2014 it had been found in 29 states (Schulz and
Tonsor 2015). PEDV is most serious in neonatal piglets where mor-
bidity and mortality can be 80 to 100%, with mortality increasing
with age. House flies have also been implicated in the transmis-
sion of hog cholera (Dorset et al.1919), transmissible gastroenter-
itis virus TGEV (Saif and Wesley 1999), and bacterial pathogens
including Lawsonia intracellularis (Dee et al. 2004, Förster et al.
2007, McOrist et al. 2011), Streptococcus suis (Enright et al. 1987,
Staats et al. 1997), and Mycobacterium (Fischer et al. 2001). House
flies can also transmit E. coli among swine causing neonatal and
post-weaning diarrhea, which are important causes of death in suck-
ling and weaned pigs respectively (Fairbrother and Gyles 2012).
Salmonella spp. can be transmitted by house flies in swine (Wang
et al. 2011) causing salmonellosis, one of the top 10 most common
diseases in weaning and grower/finisher pigs, costing pork producers
an estimated $100 million annually (Knetter et al. 2015).
While direct economic losses in swine resulting from house fly
infestations have not been documented, it has been estimated that
over $20 million is spent annually on house fly control by pork pro-
ducers in the North Central states (Campbell 1993). High numbers
10
of house flies developing in swine facilities may subject producers to
nuisance litigation. Legal cases citing fly nuisance have increased in
recent years as residential homes have increased in historically rural
areas. In 2019, over $470 million was awarded to plaintiffs in North
Carolina citing, in part, high fly numbers originating at local hog
facilities (Wall Street Journal 2018).
Sanitation practices to eliminate or minimize fly breeding materials
is the most important and effective approach to house fly management
in and around swine facilities. While there are several management
strategies for waste in hog facilities, housing on concrete slatted floors
over a slurry pit is common (USDA 2017) (Fig. 8). Alternatively, under-
floor, sloping drainage channels take liquid waste to large liquid la-
goons several times a day. Generally, the high moisture content of waste
from these management techniques prohibit house fly development.
However, both the concrete pens and waste channels are difficult to
clean thoroughly and, if not well designed, can leave areas for house fly
development. House flies are managed in swine facilities using methods
developed for dairy, beef cattle, and poultry facilities; the efficacy of
these methods in swine systems is largely unknown
Poultry
From the standpoint of fly production, poultry facilities can be div-
ided into those where: 1) birds are permitted to contact and forage
through their own feces; and 2) birds are separated above feces
which accumulates undisturbed beneath caged birds or birds held
on slatted floors. Turkeys and broiler chickens are generally housed
in wide single-story buildings (‘grow-out houses’) where birds can
move freely across a floor covered with wood shavings or other litter
material (Axtell 1986). Fly production in such facilities is low be-
cause birds scratch through the soiled litter, disturbing and drying
feces, and consuming fly immatures and other insects. In contrast,
layer hens are typically held suspended above the ground in wire
cages or aviaries, with bird feces and spilled feed accumulating in
piles or rows beneath the birds and often resulting in much greater
fly densities relative to other housing designs (Axtell 1986) (Fig. 10).
Even cage-free facilities typical for breeder birds can allow for ma-
nure to accumulate in nest boxes and under floor slats where the ma-
nure may remain undisturbed for long periods which can promote
fly development. Typically, house flies are the most abundant pest
fly in layer poultry facilities. (Lysyk and Axtell 1986a, Stafford et al.
1988, Hogsette 1993).
High fly populations are a risk to confined poultry, primarily
due to their potential to transmit pathogens among birds within
a confined setting (Shane et al. 1985). Necrotic enteritis (NE) is a
disease found worldwide wherever chickens are farmed (McDevitt
et al. 2006). This economically significant disease, caused by the bac-
terium Clostridium perfringens, results in lesions in the chicken’s in-
testine and can lead to flock mortality of 1% per day (clinical NE).
The estimated cost of NE is $2.5 billion per year in the United States
Figure 10. Manure in high-rise caged layer houses either collects below the birds or is removed by automated belt systems to liquid storage or removal off-site.
Large fans for circulation also aid in manure drying. Photos by Erika Machtinger.
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
(Wade and Keyburn 2015). Dhillon et al. (2004) found C. perfringens
in house flies on a poultry farm with both high fly populations and
NE disease incidence. Avian pathogenic strains of E. coli are also a
concern in laying hens, where they cause avian colibacillosis, sal-
pingitis/peritonitis/salpingoperitonitis (SPS) and E. coli peritonitis
syndrome (EPS). EPS is estimated to kill about 6% of the hens at
egg facilities annually, with an annual cost to the facility of about
$1.15 M (Zoetis 2018). Salmonella spp. infections are common in
poultry and result in acute and chronic diseases that consume large
economic investments for monitoring and control (Gast and Porter
2020). Flies not only disseminate Salmonella among animals, but
also can serve as a reservoir for the bacteria in the environment
(Mian et al. 2002). Naive chickens became infected with Salmonella
enteritis serovar Enteritidis when they ingested flies collected from
a facility with infected hens (Holt et al. 2007). House flies may
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also serve as mechanical vectors of viral pathogens such as turkey
coronavirus (Calibeo-Hayes et al. 2003), exotic Newcastle disease
(Chakrabarti et al. 2007, Chakrabarti et al. 2008), and avian influ-
enza (Habibi et al. 2018).
Poultry husbandry practices in intensive production systems
maximize egg production at lowest cost, but these practices can
create ideal conditions for house fly development as a result of very
high bird densities and rapid accumulation of bird feces leaving too
little time for feces to dry to prevent fly development. Conventional
caged layer farms account for over 54% of layer poultry facilities
in the United States with the majority of these (93.5%) being farms
with over 100,000 birds (USDA 2014). Manure waste is produced
at 113 g per bird per day (North and Bell 1990), which translate
to 4,129 metric tons per year for a 100,000-bird house. In many
poultry facilities, birds are housed in an artificial environment with
temperature and humidity managed in a narrow range that is quite
suitable for fly production.
There has been an increase in high-rise poultry housing from
39.7% of facilities in 1999 to 61.7% in 2013. Modern caged-layer
facilities often have automated belt systems that remove manure
daily, where it can be either placed in storage areas or taken off-site,
but such systems comprise less than 20% of facilities (USDA 2014).
Composting of stockpiled manure prevents larval development and
has the additional benefit of killing pathogens (Macklin et al. 2008).
Only 16.5% of farms store manure outside (USDA 2014).
Control of house flies is best achieved in intensive poultry sys-
tems by increased efforts toward management of feces and litter to
reduce their availability for fly production or their suitability for fly
development, primarily by using strict moisture management. When
fly outbreaks occur, most egg producers use baits and traps (77.8%)
and residual sprays for fly management (59.4%) (USDA 2014).
Over 30% use larvicides and 49.6% use space sprays or foggers.
The number of farms using biological control agents (mostly para-
sitoids) increased from 14% in 1999 to 31% in 2013. This increase
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
may be due to increased organic production, which now represents
27.6% of layer production in the United States (USDA 2014).
Dairy
Milk production since the 1980s has increased more than 59%
worldwide to 843 million tons in 2018 (FAO 2020). In developed
countries, dairy farms are growing larger (USDA 2018) and are in-
creasingly mechanized with cow management and feed carefully
controlled to increase milk production per animal. As animal density
increases, so do quantities of house fly developmental substrates.
House flies can transmit pathogens that are of concern to dairy
production, typically through mechanical methods. Although the
house fly has been implicated in transmission or transportation of
microbes that cause human illness, many of these organisms are not
pathogenic in cattle. Salmonella spp. cause substantial problems for
dairy health. Salmonellosis in cattle most commonly affects colostrum
and deficient calves can exhibit fever, diarrhea, rapid dehydration and
death within 24–48 h (Wray and Wray 2000). Both clinical outbreaks
and subclinical infections of Salmonella can drain profit from the
dairy operation by contributing to declines in milk production, abor-
tions, losses from antibiotic contaminated milk, increased culling, in-
creased labor for management of sick animals, reduced feed efficiency,
the inability to sell animals originating from an ‘infected’ herds, and
death (Holschbach and Peek 2018). Aeromonas bacteria are associ-
ated with diarrheal diseases in livestock and humans. The role of the
house flies is unclear. Flies can harbor several Aeromonas species, but
transmission studies are lacking (Nayduch et al. 2001). Within-herd
mechanical transmission of Corynebacterium pseudotuberculosis
by house flies is suspected in Israeli dairies (Yeruham et al. 1996,
Braverman et al. 1999). Yeruham et al. (2003) reported morbidity
and animal culling exceeding 6 and 16%, respectively, and milk
loss of greater than 6% in heavily affected herds. House flies have
been implicated in transmission of Cryptosporidium parvum and
Giardia lamblia, both of which cause zoonotic diarrheal diseases in
humans and livestock and have been recovered from house flies on
dairy farms (Doiz et al. 2000, Clavel et al. 2002). House flies col-
lected from dairies can be infected with Klebsiella pneumoniae, an
important cause of mastitis, however, animal management practices
are considered the primary driver of disease incidence (Nmorsi et al.
2007). Staphylococcus aureus, a cause of mastitis in dairy cattle, has
been recovered from house flies, but flies are not thought to be a
major vector of these bacteria. Recently house flies collected from
a dairy facility have been shown to carry antimicrobial- and multi-
drug-resistant coliforms, indicating that they are not only a source of
pathogenic bacteria but also harbor commensals and their resistance
genes (Neupane et al. 2020).
Figure 11. Calf housing is often a major contributor to fly production on dairy operations. Photo on left by Chris Geden; photo on right from USDA NCRS by
Scott Bauer.
11
Housing of dairy cattle can be varied. As of 2014, conventional
dairies made up 58.8% of farms in the United States and conven-
tional with grazing access farms were 26.5% of all dairies. Organic
dairy farming made up 74% of farms. Most farms either used tie
stall or stanchion (38.9%) or free stall without outdoor access
(20%) as the primary housing method (USDA 2016). Calves housed
on bedding are important sources of fly production. The use of indi-
vidual hutches to house calves is helpful for disease mitigation, but
creates substantial fly problems if hutches are not moved frequently
and provisioned with fresh bedding (Fig. 11). Bedding choice is typ-
ically straw or hay (47.1%) or sawdust (34.1%), which are very
supportive of fly larval development (Schmidtmann 1991).
In intensive dairy production systems, substantial quantities of
cattle feces collected from animal housing areas are often stored
on-site. Flushing and scraping often are used to collect manure from
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lactating cow housing areas. Although practices may vary by region,
most dairies surveyed in California use mechanical or gravity separ-
ation to sort solid manure from liquid manure (Meyer et al. 2011).
Similar to swine, recycled water from liquid manure storage ponds
or tanks is used to flush manure, while solids are most frequently
piled and remain uncovered (80.1%) or composted (26.3%) (Meyer
et al. 2011). Both solid and liquid manure are often land-applied at
some point after removal.
Intensive systems must keep substantial quantities of animal feed
on site, including hay, straw, grains, and fermenting feed additives
(Gerry 2018). Including fruit and nut waste in feed is common, much
of which is fermented either deliberately or due to placement of dry
feed in a location where it is wetted by rainfall, sprinklers, or runoff
from pens. Fermenting feed stocks can provide suitable development
substrates for house flies (Fig 12).
Cattle Feedlots
Beef production and consumption worldwide is increasing slowly with
beef consumption increasing primarily in developing countries (USDA
2020). Beef production is limited by declining rangeland availability
in most countries due to encroachment of other land uses and deg-
radation of available rangelands. Further increases in beef production
are likely to result from increasing animal density on available lands
with animals provided supplemental feeds where forage is no longer
sufficient (Bruinsma 2003). Modern cattle feedlots, where cattle do not
have access to pasture and are fed entirely on supplemental feed, are
an extreme example of beef cattle intensification (Fig. 13). The size and
animal density on U.S. feedlots continues to grow, with 40% of fed
beef cattle produced on facilities with over 32,000 animals (Economics
Research Service 2020). In April 2020, there were approximately 12 mil-
lion cattle in U.S. feedlots (National Agriculture Statistics Service 2020),
12
mostly housed at densities of 2–46 m2 of pen space per animal
(Ingvartsen and Andersen 2009, Euken et al. 2015).
As with dairy cattle, numerous human pathogens are associated
with flies from these locations, however, most are commensal or
nonpathogenic to the cattle. Large populations of flies highly cor-
related with the potential for beef cattle to shed Salmonella spp.,
indicating fly-facilitated movement of bacteria among the herd
(Vanselow et al. 2007). Neupane et al. (2019) examined the poten-
tial role of house flies in transmission of bovine respiratory disease
(BRD). Flies harbored three primary BRD pathogens of concern;
Mannheimia haemolytica, Pasteurella multocida, and Histophilus
somni were recovered from flies collected near a pen of cattle exhib-
iting BRD. Although this implicates flies as a reservoir for these mi-
crobes, the role of the house fly in the epidemiology of BRD requires
further examination.
Feedlots offer a wide range of potential house fly development
substrates of manure and animal feed. Meyer and Petersen (1983)
found that most house fly development occurred in sites that were
protected from trampling by animals, especially along fence lines
and drainage ditches. Skoda et al. (1993, 1996) found house fly
larvae to be most abundant at the interface between the feed apron
Figure 12. Stored animal feed can provide fly development sites when it
becomes wet. Photo by Chris Geden.
Figure 13. Many modern cattle feedlots pose particular problems for fly
management because of their large size and high animal densities. Source
USDA NCRS, photo by Jeff Vanuga.
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
and soil, and other studies in Nebraska (Gilbertson and Campbell
1986) and Australia (Hogsette et al. 2012) have noted that house
fly larvae are clustered along fence lines and in other protected sites
(Fig. 14). Little breeding appears to occur in manure mounds within
pens, apparently because of disturbance and trampling by the ani-
mals (Gilbertson and Campbell 1986). Protected larval development
sites along pen margins are prone to drying, and house fly popula-
tions increase after rain events (Talley et al. 2002, Hogsette et al.
2012, Urech et al. 2012). Godwin et al. (2018) modeled house fly
populations on Australian feedlots and found that rainfall events
(85–90 mm/week) resulted in elevated adult house fly populations
for 5 weeks afterward.
The large size of many modern feedlots makes it difficult to im-
plement farm-wide fly management programs and monitor their
success. Sanitation practices such as improved drainage and manure
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removal from under fence lines and other preferred development sites
are important but challenging to implement on very large operations
(Clymer 1974, Thomas et al. 1996, Talley et al. 2002). Parasitoid
releases may be helpful (Petersen et al. 1995, Floate 2003), but the
number needed to be effective on a large feedlot may be prohibitively
expensive. Parasitoid use on feedlots is discussed further below in the
section on Biological Control.
Equine
House flies can have a negative impact on the welfare of horses.
While house flies do not bite, their persistent presence on the body,
eyes, mouth, and nose of horses increase stress and their poten-
tial as disease vectors. Pigeon fever, a highly contagious condition
causing internal or external abscesses or limb infection called ul-
cerative lymphangitis, is caused by the bacteria Corynebacterium
pseudotuberculosis. House flies can transmit C. pseudotuberculosis
to horses (Barba et al. 2015). Pigeon fever was first reported in a
2015 horse health survey with 0.7% of equine farms reporting cases
(approximately 6,887 farms) (USDA 2016). House flies also are
biological vectors of Habronema spp. nematodes that can cause di-
gestive disorders, diarrhea, progressive weight loss, ulcers, colic, and
skin lesions (Amado et al. 2014, Pugh et al. 2014). Equine sarcoids,
caused by bovine papilliomavirus type 1 (BVP-1), is one of the most
common skin tumors in horses and other equids. Epidemiological
data and spontaneous development of sarcoids without direct con-
tact with affected individuals suggest flies may play an important
role as mechanical vectors of BVP and BVP-1 has been isolated from
house flies (Finlay et al. 2009).
House fly control challenges on equine facilities are generally related
to the diversity of husbandry practices, uses, and ownership. Equid
numbers on a facility may range from just one to >100 individuals.
High-value animals used for show or racing may receive very little time
in pasture, or more commonly a few hours to half a day. Conversely,
pleasure or ranch horses may be pastured individually or in groups with
little stall time. Bedding material used in stalls varies from straw, wood
shavings, sawdust, or even newer products containing paper, peanut
hulls, or hemp. Forage and feeding may be primarily from pasture or
can be supplemented with hay fed in small or large flakes, or in large
hay round bales as with cattle. Manure management practices vary
with availability of land and local or regional regulations. Most facil-
ities use manure pits or piles. Piles can be removed at regular intervals,
left to accumulate, or applied to pastures or crop fields (USDA 2016).
Along with region and local conditions, management choices can
influence house fly presence. House flies may develop in manure ac-
cumulation areas, pasture sheds or animal aggregation areas, waste
hay in fields, or in stalls. In choice tests, house flies developed better
in equine manure alone and manure mixed with pine shavings than
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
Figure 14. Most fly larval development in feedlots occurs in protected sites such as fence lines and gaps between and beneath feeders, where spilled feed gets
mixed with manure. Photos by Chris Geden.
in manure mixed with hay or straw (Machtinger et al. 2014), sug-
gesting bedding choice influences fly development as it does in calf
hutches (Schmidtmann 1991). In well-managed facilities, high house
fly numbers may be a result of emigration from local or neighboring
livestock or poultry facilities.
To reduce house fly presence on horses, many owners and man-
agers rely on commercial fly repellents. In 2015, 76% of equine
operations used on-animal repellents (USDA 2016), and 36.8% re-
ported applying insecticides in or near equine housing areas, usu-
ally in the form of automatic misters. However, most commercial fly
repellents and residual spray products are pyrethroid-based. These
products generally have low concentrations of pyrethroid because of
equine dermal sensitivity to higher levels (Stevens et al. 1988). High
levels of pyrethroid resistance in house fly populations throughout
the country have reduced the residual activity of many of these prod-
ucts to less than a few hours (Tuorinsky and Machtinger 2020) and
thus they may not offer the protection needed to reduce annoyance
or pathogen transmission. Recently, product formulations including
fatty acids, geraniol, and other alternatives have been shown to in-
duce more behavioral inhibition of house flies in laboratory settings
and have longer residual activity (Tuorinsky and Machtinger 2020).
Monitoring and Management
House fly’s short development time, high fecundity, mobility, ability
to exploit a myriad of developmental substrates, adaptability, and
notorious propensity for developing resistance to new insecticides
combine to make it challenging to manage. Maintenance of fly popu-
lations below acceptable levels requires a diligent IPM approach that
takes full advantage of the many opportunities to combine moni-
toring with cultural, mechanical, biological, and chemical control
approaches. Examples of available and potential tools for each of
these elements is presented in the following sections.
Monitoring
Methods to monitor house fly activity have been described and
evaluated for poultry facilities (Anderson and Poorbaugh 1964,
Axtell 1970, Burg and Axtell 1984, Beck and Turner 1985, Lysyk
and Axtell 1986), dairies (Pickens et al. 1972, Pickens and Miller
1987, Gerry et al. 2011), and beef cattle (Urech et al. 2004). General
reviews of house fly monitoring methods and applications of those
methods are also available (Lysyk and Moon 1994; Gerry 2020).
In 1945, Harvey Scudder was assigned by the U.S. Public Health
Service to assess the efficacy of DDT for house fly control. To do
13
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Figure 15. The Scudder grid can be useful for making instantaneous fly
counts when only one species is present. Photo by Chris Geden.
this, he developed what is now known as the Scudder grid sampling
method (Scudder 1947, 1996). The grid, a framed series of parallel
wooden slats, is thrown to the ground where flies are concentrated
and counts are made after the flies have settled again, usually 30 s
later (Fig. 15). When multiple counts are made at several locations
at a field site, the method can provide a simple and inexpensive in-
stantaneous index of fly abundance at a location (Dhillon and Chalet
1985, Murvosh and Thaggard 1966). Population changes can also
be monitored by repeated sampling over time (Madwar and Zahar
1951). Because house fly activity varies considerably with time of
day and changing environmental conditions (Parker 1962, Zahn and
Gerry 2020), grid counts should be recorded at a consistent time of
day and with similar environmental conditions when sites are to be
visited on multiple occasions.
Sticky traps of various kinds also can be used to monitor house
flies. The conventional fly ribbon (Fig. 16) has been used for many
years and provides more reliable estimates of fly abundance than grid
counts (Anderson and Poorbaugh 1964, Raybould 1966, Pickens
et al. 1972, Nurita et al. 2008). Ribbons also have the advantage
of providing time-lapse rather than instantaneous counts and allow
data collection on other species when present. Disadvantages of
sticky ribbons are dust accumulations that compromise the adhe-
sive and underestimations at high fly densities when tapes reach
their carrying capacity of flies before they are collected (Rutz and
Axtell 1981, Kaufmann et al. 2001a, Gerry et al. 2011). A variation
on stationary tapes is the ‘walking sticky tape’ method, where a
14
Figure 16. Sticky tapes and cards are commonly used for monitoring house flies, especially when species identification is needed. Photos by Erika Machtinger.
Figure 17. Baited jug trap used for monitoring house flies in indoor settings.
This method is only effective when flies are sufficiently susceptible to the bait
toxicant to die immediately after feeding. Photo by Chris Geden.
person carries the tape and walks at a consistent pace. This method
has the advantage of providing an immediate estimate of fly abun-
dance and has mostly been used to survey the upstairs of high-rise
poultry houses (Turner and Ruszler 1989, Hinton and Moon 2003,
Kaufman et al. 2005b). Peel-off sticky cards are a convenient alter-
native to ribbons that are easy to place and transport (Hogsette et al.
1993, Bell et al. 2019).
Burg and Axtell (1984) described the use of baited jug traps for
monitoring indoor populations of flies. These traps were standard
1-gallon milk jugs with holes cut in their sides that were baited with
a dry sugar-insecticide bait and suspended from ceilings (Fig. 17).
Flies entered the traps, fed, and died in place. The trap was used for
several years (Lysyk and Axtell 1985, 1986a; Beck and Turner 1985;
Stafford et al. 1988), but fly resistance to the toxicant in the bait
(methomyl) rendered it impractical until new fast-killing insecticides
became available. The method may be useful in locations where
neonicotinoids are still effective. However, the use of this method for
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Figure 18. Spot cards are highly effective for monitoring relative fly activity
over time if only one species is present. Cards with printed grid lines can
help when making spot counts. Counting spots is challenging when fly
populations are high. Photos by Erika Machtinger.
long-term monitoring (over years) must be carefully considered as
increasing insecticide resistance in the fly population will reduce trap
counts unrelated to fly density or activity.
Spot cards (Fig. 18) are the most widely used monitoring
method. First described by Axtell (1970), spot cards are standard
7.62 × 12.70 cm (3 × 5 inch) white cards that are attached to
building structures and left in place to accumulate fly fecal and
regurgitation spots. Cards are typically left in place for a week.
A number of studies have compared spot cards with other sam-
pling methods, and attempts have been made to use the cards to
estimate absolute fly numbers (Beck and Turner 1985; Lysyk and
Axtell 1985, 1986; Gerry et al. 2011). Although card counts can
vary widely depending on temperature and card location and the
fly species cannot be distinguished, they are easy and inexpensive
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
to use and can be stored for years. They are particularly effective
for monitoring relative abundance of indoor fly populations over
time. In total, 100 spots per card per week has long been cited as
an action or nuisance threshold (Axtell 1970), but this threshold
is not based on scientific evidence.
Most of the work discussed above has concentrated on rela-
tively enclosed systems such as poultry houses and dairy barns in
cooler climatic zones. These systems have geometric consistency and
constraints that are helpful when implementing a monitoring plan.
Monitoring is more complicated when a large portion of the fly
population occurs outdoors such as on equine farms, feedlots, and
large dairy farms in warmer parts of the world. Gerry et al. (2011)
reported that spot cards were the most effective sampling method
at dairies in Southern California, where frequent rains are not a
problem. They also described a software application (FlySpotter)
that automates the tedious process of counting the numbers of
spots on cards (available at https://www.veterinaryentomology.org/
flyspotter-house-fly-monitoring). Where frequent rains preclude the
practical use of spot cards, sticky traps and liquid jar traps can be
effective monitoring tools for outdoor fly populations. (Geden 2005,
2006; Gerry et al. 2011; Urech et al. 2012; Aziz et al. 2016). Baited
plastic strips with toxicant can also be placed in sheltered locations
for monitoring (Geden 2005), but only if flies are sufficiently suscep-
tible to die close to the strips.
Cultural Control
Manure management is the cornerstone of any fly control program.
Manure removal schedules are particularly critical. Mature larvae
begin searching for pupation sites after 4–6 d under warm condi-
tions, and often pupate away from the manure in protected and dry
areas that are not cleaned out. Manure removal in facilities other
than layer poultry therefore needs to be done at least twice a week to
be effective during summer months. In caged-layer poultry systems,
leaving residues of old manure during house cleanouts would be ex-
pected to encourage colonization of fresh droppings by fly predators,
but recolonization by predators is slow under alternate row removal
schedules (Peck and Anderson 1070, Geden and Stoffolano 1988;
Mullens et al. 1996b, 2001). Leaving a pad of old manure may pro-
mote drying of fresh deposits and predator colonization (Hinton and
Moon 2003), but Mullens et al. (1996a) observed no benefit from
this practice in open-sided houses in California. Frequent removal of
solid manure to liquid storage systems also prevents fly larval devel-
opment, but such systems can produce troublesome populations of
biting midges (Culicoides) and Culex mosquitoes.
When manure is removed, house fly larvae that are present in
that manure can complete development after it is applied to crop-
land, even when it is incorporated into or covered with soil. Watson
et al. (1998) estimated 89% of larvae in infested poultry manure
were killed by mechanical damage during application with a manure
spreader, and that only 25% of larvae produced adult flies when they
were covered with 30 cm of soil. In spite of this attrition, the authors
estimated that over 300,000 flies could emerge from a field on which
manure from a poultry house had been applied. Tahir and Ahmad
(2013) observed some fly emergence when pupae were buried under
30 cm of heavy clay soil, and Cook et al. (2020) found that house
fly emergence from soil was unaffected by severe compaction.
Composting is an alternative that can render manure unsuitable for
fly larval development (Moon et al. 2001, Abu-Rayyan et al. 2010).
Moisture mitigation is another important component of cultural
control. Because house fly larval development is optimal in substrates
with 50–75% moisture (Fatchurochim et al. 1989), tactics that aid
15
in drying out fly larval habitats are beneficial for managing fly popu-
lations. Water management practices include maintaining waterers,
addressing drainage problems, and improving airflow in enclosed
systems through the use of tunnel or turbo ventilation. Harrowing
manure in animal pens can also aid in drying.
Mechanical and Physical Control
Mechanical and physical control options kill pests directly or make
the environment unsuitable or inaccessible for pest flies. Screens or
air curtains can reduce the entry of flies into sensitive areas such as
milk and egg rooms (Mathis et al. 1970, Carlson et al. 2006), and
high-speed fans in poultry houses can affect fly distribution in ma-
nure pits (Geden et al. 1999). Additives like hydrated lime, Ca(OH)2,
acetic and boric acid (Lachance et al. 2017) and sodium bisulfate
(Sweeny et al. 2000, Calvo et al. 2010) have been used to dry ma-
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nure and to make the pH of manure less suitable for fly development.
Ultraviolet light has long been recognized as a house fly attractant
(Pickens et al. 1969, Pickens and Thimijan 1986). Electrocuting
light traps sometimes have been reported to kill enormous num-
bers of flies in poultry houses (Rutz et al. 1988, Pickens et al. 1994,
Hogsette 2019), but there are no data indicating that they are ef-
fective at reducing populations. UV traps are useful against small fly
populations in sensitive areas such as dining areas and grocery stores
(Lillie and Goddard 1987). Traps with glue boards are preferred to
electrocuting models, as electrocution can result in dissemination of
fly-borne pathogens into the local air space (Tesch and Goodman
1995, Urban and Broce 2000).
Sticky ribbons, sheets and cards can be useful but also are limited
by the number of flies they can collect before becoming saturated
and by the adhesive becoming ineffective from dust. Giant sticky rib-
bons are available in rolls that greatly expand the available surface
area of adhesive traps. Kaufman et al. (2001a) reported that such
large ribbons collected over 9 million flies during a 10-wk period in
a poultry house and resulted in a measurable reduction in fly abun-
dance in the upstairs of the house. In a subsequent study (Kaufman
et al. 2005a), large sticky ribbons collected 900,000 house flies in a
calf barn over 10 wk.
Odor-Based Traps and Baits
Attractant-based traps are a mainstay of fly management. There is a
vast literature on house fly attractant strategies. Harper (1872) re-
ceived a patent for the first inverted cone trap, and Howard (1911)
described a bait made with fish heads, watermelon rinds, corncobs,
and ice cream. Much of the research on fly attractants has focused on
identifying components of food odors that can be incorporated into
lures (Garrett 1965, Frishman and Matthysse 1966, Mayer 1971,
Mulla et al. 1978). Early efforts with baits relied on natural prod-
ucts such as fermented egg slurries (Willson and Mulla 1973) or
combinations of items such as molasses, milk, yeast, grain, blood,
and banana extract (Pickens et al. 1973, Pickens and Miller 1987).
Brown et al. (1961) tested a range of defined chemical attractant
candidates and found that combinations were superior to any in-
dividual component tested alone. Mulla et al. (1977) reported that
blends of trimethylamine, ammonia, indole, and linoleic acid were as
attractive to house flies as natural food baits. More recently, Hung
et al. (2015) found that house flies were attracted to honeydew-
contaminated plant material and that associated fungi may play a
role in the attraction. Specific compounds associated with honeydew
contaminated plants that were attractive to house flies were (Z)-3-
hexenyl acetate and benzaldehyde (Hung et al. 2019). Tang et al.
(2016) examined components of fermenting wheat bran and found
16
that a blend of ethyl palmitate, ethyl linoleate, methyl linoleate, and
linoleic acid was attractive to gravid females.
In addition to feeding attractants, flies are attracted to (Z)-
9-tricosene (muscalure) (Carlson et al. 1971, Carlson and Beroza
1973). The most common commercial feeding-attractant in use
today, the Farnam Fly Attractant (now branded as the Starbar Fly
Trap Attractant by Central Life Sciences), derived by modifying
ratios of trimethylamine and indole and adding (Z)-9-tricosene. This
attractant is used in granular sugar baits with a toxicant and in liquid
jar traps without a toxicant such as the Terminator and Captivator
brands. A wide variety of attractant-baited traps without toxicants
are now available on the market, most of which are variations on the
‘jar’ or ‘jug’ design that lure flies into a container containing liquids
in which the dead flies accumulate (Geden et al. 2009). Because these
types of traps rapidly become ‘saturated’ when fly populations are
high (Fig. 19), require frequent maintenance, and are malodorous,
they are most appropriate for outdoor environments with light to
moderate fly populations.
Biological Control
Many methods have been researched for biological control of house
flies including fungal, bacterial, or viral pathogens as well as pred-
ators, parasitoid wasps, and parasitic nematodes. Many of these
methods have been explored extensively, while others are relatively
new or have yet to be developed fully. There are specific benefits and
challenges associated with each method that should be considered
prior to incorporation in an IPM plan.
Entomopathogenic Fungi
Entomopathogenic fungi are naturally occurring fungi that kill in-
sects and closely related arthropod hosts. They enter the host by
invading the cuticle or being ingested, where they quickly proliferate,
disseminating hyphal bodies and releasing various toxins. After the
death of the host the fungus may sporulate on the exterior of the
host cadaver if the humidity is high enough (Charnley 1989, Samson
et al. 1988). Conidia are then dispersed to other uninfected individ-
uals and the cycle continues (Kurtti and Keyhani 2008). The primary
entomopathogenic fungi that have been evaluated for house fly con-
trol are Metarhizium brunneum (=anisopliae), Beavueria bassiana
(Fig. 20), and Entomophthora muscae.
Metarhizium brunneum Petch and B. bassiana Balsamo-Crivelli,
while taxonomically distinct, are often tested and treated similarly
in the literature. These are soil-dwelling fungal pathogens that are
Figure 19. Jar traps with liquid attractants and no toxicant are a mainstay of fly management but can fill with flies quickly, require frequent servicing, and are
malodorous. Photos by Chris Geden (left) and Erika Machtinger (right).
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
naturally found in filth fly populations. It is important to note that
the M. anisopliae lineage was revised in 2009 (Bischoff et al. 2009).
Many isolates of M. brunneum have been researched for biocon-
trol with some strains being developed as mycopesticides, however
many of the articles detailing this research were published prior to
2009, when these isolates were classified as M. anisopliae. Here we
refer to all M. anisopliae/brunneum isolates as M. anisopliae Sensu
lato (s.l.).
The potential of B. bassiana as a biological control agent against
pest flies was suggested by Dresner (1950). Steinkraus et al. (1990)
first reported B. bassiana infections at a low prevalence (~1%) in
house flies from New York state dairy farms. Since then, an abun-
dance of literature has documented the potential efficacy of these
fungal species to infect and kill immature and adult filth flies under
laboratory (Hall et al. 1972, Kuramoto and Shimazu 1992, Barson
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et al. 1994, Geden et al. 1995, Watson et al. 1995, Angel-Sahagún
et al. 2005, Lecuona et al. 2005, Kaufman et al. 2008, Weeks et al.
2017), and field conditions (Watson et al. 1996, Kaufman et al.
2005b, Cova and Scorza-Dagert 2006).
Beauveria bassiana and M. anisopliae s.l. have been tested
against flies of various life stages, ages, and physiological character-
istics. While house fly age does not seem to affect pathogenicity of
B. bassiana or M. anisopliae s.l. (Rizzo 1977, Kaufman et al. 2008),
female flies seem to be more susceptible than males (Rockstein
and Lieberman, 1958, Lecuona et al. 2005, Anderson et al. 2013,
Acharya et al. 2015a), and infection also reduces reproductive fitness
(Acharya et al. 2015a). Horizontal transfer of conidia from males to
females may be an avenue of dissemination of fungi in an otherwise
adverse environment for fungal dispersal (Cárcamo et al. 2015).
Entomopathogenic fungi effects on immature house flies have
been variable. Machtinger et al. (2016a) found decreased eclosion
rates of house fly eggs treated with some commercial formulations of
B. bassiana and M. anisopliae s.l. Exposure of larvae to B. bassiana
has had little effect in some studies (Geden et al. 1995, Lecuona et al.
2005) and caused higher mortality in others (Steinkraus et al. 1990,
Barson et al. 1994, Darwish and Zayed 2002). Young larvae are more
susceptible than older larvae (White et al. 2020), possibly due to cu-
ticular properties of developing larvae (Boucias and Pendland 1991).
Immature flies may be more susceptible to M. anisopliae s.l. than
B. bassiana (Bernardi et al. 2006, Fernandes et al. 2013). The responses
of immature filth flies to fungal infection may also depend on the viru-
lence of the fungal isolate, assay method, fly colony variability, conidial
doses, substrate used, and culture methods. Larval resistance to fungal
infection may be related to the strong immunological defenses that flies
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
Figure 20. House fly cadavers with Beauveria bassiana conidial blooms.
Photo by Chris Geden.
need to cope with the diversity of microorganisms found in their devel-
opmental environment (Nayduch and Joyner 2013). House fly larvae
provide few suitable attachment sites for conidia and their constant
movement through the larval substrate provides ample opportunity for
conidia to be dislodged.
Various application methods for B. bassiana and M. anisopliae
s.l. have been examined in laboratory and limited field settings,
including topical applications (Sharififard et al. 2011), baits (Geden
et al. 1995, Watson et al. 1995, Renn et al. 1999, Lecuona et al.
2005, Hong and Hai 2012, Mishra et al. 2013, Machtinger et al.
2016c), residual sprays (Watson et al. 1995, Blanford et al. 2012,
Acharya et al. 2015b), and dusts (Geden et al. 1995, Watson et al.
1995). Aqueous and dust formulations can be applied directly on
target flies or as a spray into the environment. Use of baits also has
been suggested to slow insecticide resistance, reduce control costs,
and reduce the effects on nontarget organisms by targeting fungal
exposure to flies (Vega et al. 1995, Zimmer et al. 2010). However,
there have been no published field studies assessing the efficacy
of baits containing M. annisoliae s.l. or B. bassiana for control of
house flies.
Significant differences in virulence have been noted among iso-
lates of B. bassiana and M. anisopliae s.l. Fungal adaptation to host
or host to fungal enzymes may influence virulence (St. Leger et al.
1986). Isolate differences have been noted in house fly adult and larval
mortality (Mwamburi et al. 2010, Sharififard et al. 2011, Mishra and
Malik 2012). When isolates adapt to their current host they may lose
plasticity for invading other arthropods. Thus, the search for fungi
to be used as control agents against filth flies should include isolates
from the target insect (Poprawski et al. 1985, Steinkraus et al. 1991).
Potential synergism between entomopathogenic fungi and bacterial
pathogens or insecticides (Mwamburi et al. 2009, Sharififard et al.
2011, Farooq and Freed 2016, Johnson et al. 2018) may provide
additional opportunities to increase virulence of fungi.
Virulence of B. bassiana and M. anisopliae s.l. to house flies
may depend on environmental conditions. Temperature and relative
humidity affect germination of conidia or virulence (Sivasankaran
et al. 1998, Arthurs and Thomas 2001, Sharififard et al. 2012,
Mishra et al. 2015). Ultraviolet light exposure, dust and particu-
late debris, and ammonia from livestock facilities can affect effi-
cacy of M. anisopliae s.l. and B. bassiana (Leland and Behle 2005,
Watson et al. 1995, Acharya et al. 2015b). These are serious consid-
erations for development and deployment of M. anisopliae s.l. and
17
B. bassiana in livestock, poultry, or equine facilities, but fungi may
be protected by appropriate formulations (Morley-Davis et al. 1996,
Alves et al. 1998, Reis et al. 2008).
Relatively few field studies have been conducted evaluating
entomopathogenic fungi as filth fly control agents. An integrated
management program including a spray formulation of B. bassiana
in conjunction with pupal parasitoid releases was successful in
caged-layer poultry in New York state (Kaufman et al. 2005b).
The number of fly larvae recovered from the treated facilities was
less than half the number recovered from pyrethrin treated facil-
ities, moreover there was a greater recovery of beneficial insects.
Watson et al. (1996) observed reduced adult house fly numbers
after spray applications of B. bassiana in calf hutches. Applications
of M. anisopliae s.l. reduced populations of house flies in treated
poultry sheds two-fold compared to the control sheds (Fernandes
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et al. 2013). Conversely, weekly applications of B. bassiana were
inadequate to achieve effective larval control in poultry houses in
South Africa (Mwamburi et al. 2009).
The Entomophthora muscae (Cohn) Fresen (Entomophthorales:
Entomophthoraceae) species complex, which includes E. muscae
and E. schizophorae Keller and Wilding (Pinnock and Mullens
2007), is a group of fungi that have generally larger and more fragile
conidia, faster germination times, a stronger tendency to produce
conspicuous epizootics, and a narrower host range than the group
that includes M. anisopliae s.l. and B. bassiana (Pell et al. 2001).
Transmission of E. muscae occurs during a characteristic behavior
change that causes house flies to anchor themselves to an elevated
structure in the environment (Krasnoff et al. 1995). Following the
death of the fly, large and sticky conidia are discharged into the en-
vironment where they can be encountered by healthy flies (Mullens
and Rodriguez 1985, Six and Mullens 1996a, Kalsbeek et al. 2001a).
Male flies, attracted to swollen, infected cadavers, can transfer co-
nidia to healthy females (Watson and Petersen 1993, Zurek et al.
2002) and the infection renders females sterile.
Natural epizootic infections of E. muscae are well documented
and primarily occur in the fall when temperatures are moderate
and populations of flies are still high (Mullens et al. 1987a, Watson
and Petersen 1993, Steinkraus et al. 1993, Six and Mullens 1996b).
Because of the temperature sensitivity of E. muscae, flies are able to
induce “behavioral fever,” whereby infected individuals locate areas
of higher temperature to elevate their body temperature and kill the
infection (Watson et al. 1993, Kalsbeek et al. 2001b). To what extent
natural populations perform this behavior is not known.
Use of E. muscae for house fly control has significant challenges.
The conidia of E. muscae are fragile and cannot be held for future
use like M. anisopliae s.l. and B. bassiana (Kalsbeek et al. 2001a).
Methods for mass production of E. muscae have been developed
(Kramer and Steinkraus 1981, Mullens 1986), but the introduction
of the fungus to the target population requires using either live in-
fected flies or cadavers of recently infected flies. Attempts to control
house fly populations with E. muscae in the field have had limited
success to date (Steinkraus et al. 1993, Geden et al. 1993, Six and
Mullens 1996b).
Entomopathogenic Bacteria
Bacillus thuringiensis Berliner (Bt) is an aerobic, Gram-positive,
spore-forming bacterium (Mwamburi et al. 2009). Bacillus
thuringiensis can be isolated from many environmental sources
such as soil, insects, and coniferous and deciduous leaves (Schnepf
et al. 1998). Formulations of Bt have been used for decades as
biopesticides for many agricultural pests.
18
Early work demonstrated that feeding animals (cattle and
poultry) Bt could control larvae developing in the resulting ma-
nure (Burns et al. 1961, Miller et al. 1971). Larval control was also
obtained by mixing Bt into larval development sites (Rupes et al.
1987). However, fly resistance to the exotoxins produced by the
tested Bt strains quickly made these techniques unsuitable for con-
trol (Harvey and Howell 1965, Wilson and Burns 1968). Additional
vertebrate safety concerns were raised, which led to the prohibition
of exotoxin-producing Bt in the United States (McClintock et al.
1995, Tsai et al. 2003).
Subsequent evaluations of exotoxin-free Bt were disappointing.
However, several strains of Bt were identified that were effective
against house fly larvae and adults (Indrasith et al. 1992, Hodgman
et al. 1993, Johnson et al. 1998, Zhong et al. 2000) and it was dis-
covered that the δ-endotoxin Cry1B was found in all the Bt strains
that were active against house flies. The endotoxins are safe for hu-
mans, other vertebrates, plants, and are biodegradable (de Barjac
1978, Adang et al. 2014).
Current research has focused on Bt for house fly control in topical
applications and as a feed additive. While application of Bt to larval
development sites has resulted in some mortality of flies, greater suc-
cess has been from use of Bt as an animal feed additive. Promising
control when fed to poultry (Labib and Rady 2001; Mwamburi
et al. 2009, 2011; Merdan 2012) and horses (Martins 2013) has
been observed. There is promise in the use of Bt as a biological con-
trol method for house flies in a variety of animal production systems.
Salivary Gland Hypertrophy Virus
House fly salivary gland hypertrophy virus (MdSGHV) is a house
fly-specific virus that replicates in salivary glands, generally resulting
in obvious hypertrophy of the glands and sterility of female hosts
(Lietze et al. 2011b) (Fig. 21), although virus particles can be found
in other tissues (Lietze et al. 2011a, Kariithi et al. 2017). House flies
are infected with MdSGHV globally, with prevalence rates usually
of less than 2% (Geden et al. 2008, 2011a; Prompiboon et al. 2010;
Lietze et al. 2011b; Lietze et al. 2013).
Infection is thought to occur via food sources shared between in-
fected and healthy flies (Lietze et al. 2009, 2013); thus, infective baits
have been considered for fly control purposes. However, feeding on
infected material results in limited infection of flies in the laboratory
(Geden et al. 2008, 2011a; Lietze et al. 2009; Lietze et al. 2011c),
likely because house fly susceptibility to the virus is only during a
brief window after adult eclosion, when they would not normally be
feeding. The PM is an effective barrier to oral infection, as suscepti-
bility in older flies can be restored if they are given drugs that disrupt
the PM (Boucias et al. 2015). An alternative approach to infective
baits is the use of a homogenate of infected flies sprayed in areas of
house fly activity (Geden et al. 2011). However, the efficacy of this
approach has not been demonstrated in the field.
Predators
Three families of mites, Macrochelidae, Uropodidae, and
Parasitidae, are known to prey upon fly eggs or larvae and nat-
urally occur in poultry manure (Axtell 1961, 1963a). One of the
most common predatory mites is Macrocheles muscaedomesticae
(Scopoli) (Fig. 22). Some experiments have demonstrated upwards
of 20 house fly eggs consumed by M. muscaedomesticae per mite
per day (Geden and Axtell 1988, Geden et al. 1988). Reductions
in house fly numbers in cattle manure and poultry manure have
been demonstrated under simulated field conditions with the use
of M. muscaedomesticae (Axtell 1986). Reviews of the literature
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
and an assessment of Macrochelidae as biological control agents
have been published (Axtell 1969, Geden 1990). Populations of
M. muscaedomesticae are important for the natural suppression of
house flies (Axtell 1963b), however this species has not been devel-
oped commercially.
Many beetles are important predators on fly eggs and larvae,
but the major predaceous species is the histerid Carcinops pumilio
(Erichson) which often is very abundant in poultry manure; adults
and larval stages prey on fly eggs and larvae (Morgan et al. 1983)
(Fig. 22). A single beetle adult may consume 13–83 house fly eggs
per day and beetle larvae consume 13–26 eggs per day (Geden and
Axtell 1988, Geden et al. 1988). The wide distribution of these bee-
tles as well as abundance and high rate of predation make C. pumilio
a major predator of house flies. These beetles are typically found
in poultry pit houses (Geden 1984) and are less common in other
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animal facilities. Carcinops pumilio can be purchased from commer-
cial insectaries for release, and populations can be collected with
species-specific traps to re-release after pit cleanout.
Other fly species are also predators of immature house flies. Black
dump fly (Hydrotea aenescens [Wiedemann]) is a naturally occurring
muscid that is most commonly found on poultry and swine facilities.
It is known in the older literature as Ophyra aenesens Weideman.
Larvae of this species are facultative predators that can develop in
substrates without prey (Hogsette and Washington 1995, Farkas
and Hogsette 1998, Hogsette et al. 2002) but are also capable of
killing 15 house fly larvae per day (Geden et al. 1988). Adult flies
prefer dark spaces and are less prone to becoming nuisance pests
than house flies (Nolan and Kissam 1987). Black dump flies have
been studied as potential biological control agents for house flies,
and augmentative releases sometimes have resulted in its dominance
over house flies (Nolan and Kissam 1985, Turner and Carter 1990,
Turner et al. 1992). They are available from commercial insectaries
in the United States and Europe. However, there are concerns about
release of this species as their ability to transmit pathogens mechan-
ically may be similar to that of other pest flies (Olsen and Hammack
2000, Szalanski et al. 2004).
Parasitoids
Naturally occurring pupal parasitoids (Hymenoptera: Pteromalidae)
have been used for control of house flies on animal facilities for
nearly 50 yr. House fly parasitoids occur naturally in all areas where
suitable hosts can be found. Naturally occurring parasitoid popula-
tions are not sufficient to manage flies at acceptable levels because
they are slower to develop than the fly host. However, augmenta-
tive releases of parasitoids can be effective at suppressing pest fly
populations when coupled with other management methods (Geden
et al. 1992, Skovgård and Nachman 2004, McKay et al. 2007).
Parasitoids are commercially available, and their use has increased
by some commodity groups, likely in part due to the easy availability
(USDA 2006, Machtinger et al. 2013). The taxonomy, natural his-
tory, and use of parasitoids have been reviewed in Machtinger and
Geden (2018) and Machtinger et al. (2015b). Here we will review
available species for house fly control programs and use on the target
animal commodity groups.
Although a wide diversity of species parasitize house fly pupae,
the most commonly collected and commercially available parasit-
oids include Muscidifurax raptor Girault and Sanders, M. zaraptor
Kogan and Legner, M. raptorellus Kogan and Legner, Spalangia
cameroni Perkins, S. endius Walker, Nasonia vitripennis (Walker),
and Trichomalopsis sarcophagiae (Gahan) (Fig. 23). Most of these
species have similar life histories where the female parasitoid
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
locates a suitable fly puparium, drills through the puparium, and
deposits either one (solitary species) or multiple eggs (gregarious
species) on the surface of the host pupa (Gerling and Legner 1968).
The resulting parasitoid larva (or larvae) consumes the pupa and
emerges as an adult 2–4 wk later. Development of a parasitoid
from egg to adult takes 14–30 d under warm conditions depending
on species, sex, host, temperature, environment, and biotypes
(Birkemoe et al. 2012).
Figure 21. A healthy fly (A) and a fly infected with salivary gland hypertrophy
virus (B), showing underdeveloped ovaries (Ov) and overdeveloped salivary
glands (Sg) in the infected fly (Mg = midgut). Photo by Lyle Buss, University
of Florida (Florida, USA).
Figure 22. The two most important predators of house fly immatures are the mite Macrocheles muscaedomesticae (left) and the histerid Carcinops pumilio.
Photos by Eric Palevsky, Agricultural Research Organization, Israel and Erika Machtinger, respectively.
19
Evaluations of parasitoid releases on filth fly populations
have demonstrated reduction in some situations (Morgan and
Patterson 1990, Geden et al. 1992, Petersen and Cawthra 1995,
Crespo et al. 1998, 2002, Skovgård and Nachman 2004, Geden
and Hogsette 2006), but not in others (Morgan 1980, Meyer et al.
1990b, Andress and Campbell 1994, Weinzierl and Jones 1998,
McKay and Galloway 1999, Kaufman et al. 2001c). Biological
characteristics of released parasitoid species that may influence ef-
fectiveness include parasitoid microhabitat preferences, intra- and
interspecific competition (Machtinger and Geden 2015, Taylor
et al. 2016), and factors influencing parasitoid abundance and dis-
tribution (Skovgård 2004). Colony quality, previously established
parasitoid populations, and the lack of understanding of timing
and methods of parasitoid release could also have impacted the
success of releases (Peterson et al. 1983, Patterson and Rutz 1986,
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Legner et al. 1990). Nasonia vitrpennis does not appear to be ef-
fective in poultry, dairy, cattle feedlot, or swine systems (Stage and
Petersen 1981, Meyer et al. 1990b, Andress and Campbell 1994,
McKay and Galloway 1999, Kaufman et al. 2001c, Birkemoe
et al. 2004).
Parasitoid releases in poultry facilities were some of the earliest
evaluations (Legner and Brydon 1966). The relative stability of
habitat in poultry facilities and presence of other predators, including
beetle and mites, make these facilities well-suited for parasitoid re-
leases (Geden 1990). Reduction in house fly numbers has been
achieved with releases of S. endius (Morgan et al. 1975a; Morgan
et al. 1975b; Morgan et al. 1981a,b; Morgan and Patterson 1990),
M. raptor and M. raptorellus (individually or in tandem) (Rutz and
Axtell 1979, Crespo et al. 1998, Kaufman et al. 2002, Al-Ani et al.
2012), and combinations of M. raptorellus and S. cameroni (Geden
and Hogsette 2006, McKay et al. 2007).
Dairy facilities often have fly-breeding sites that have some
measure of habitat constancy and environmental protection.
However, house fly control as a result of parasitoid releases has
been more variable. High levels of parasitism have been achieved
with releases of S. endius (Morgan and Patterson 1977), S. cameroni
(Skovgård 2004; Skovgård and Nachman 2004), M. raptor
20
Figure 23. Examples of the two most important genera of house fly parasitoids; Spalangia (S. cameroni, left) and Muscidifurax (M. raptor, right). Photos by
Erika Machtinger.
(Geden et al. 1992), and M. raptor and M. raptorellus combinations
(Kaufman et al. 2012). Conversely, M. zaraptor and M. raptor re-
leases had negligible impacts on adult fly populations on dairies
(Meyer et al. 1990b, Miller et al. 1993), although numbers of para-
sitoids released were not as high as in other studies.
In cattle feedlots, fly development sites are spread over large
areas and often disturbed and exposed to the environment, making
parasitoid-based fly control more challenging. The range of dis-
persal for pteromalid parasitoids is limited (Pawson and Petersen
1988, Tobin and Pitts 1999, Skovgård 2002, Machtinger et al.
2015c). Releases of S. endius (Morgan 1980), and locally collected
M. zaraptor and M. raptorellus (Petersen et al. 1992, 1995; Petersen
and Cawthra 1995; Petersen and Curry 1996), S. nigroaenea Curtis
(Weinzierl and Jones 1998) and combinations of M. raptorellus and
Trichomalopsis sarcophagae Gahan (Floate 2003, Floate et al. 2000)
have been effective to some degree in reducing fly populations in
confined cattle. However, S. endius releases (Petersen et al. 1983)
and mixed releases of M. zaraptor, N. vitripennis, S. endius, and
S. nigroaenea (Andress and Campbell 1994) have not demonstrated
control. In both these latter cases, species may have been released in
areas where they were not well-adapted.
Information on use of parasitoids at other animal facilities is
limited. On swine facilities, releases of S. endius and S. cameroni
have had some effect (Morgan 1980, Skovgård and Nachman 2004).
While Spalangia spp. have been the dominant species recovered from
equine farms (Pitzer et al. 2011, Machtinger et al. 2016b), these
studies were limited to surveillance.
Parasitic Nematodes
Entomopathogenic nematodes in the families Steinernematidae
(Steinernema spp.) and Heterorhabditidae (Heterorhabditis spp.)
(Order: Rhabditida) have been reviewed extensively (Kaya and
Gaugler 1993, Gaugler 2002, Georgis et al. 2006, Koppenhöfer 2007,
Poinar and Grewal 2012, Atwa 2014). These parasites are vectors of
bacterial symbionts in the genera Xenorhabdus and Photorhabdus
(Enterobacteriales: Enterobacteriaceae), respectively (Forst et al.
1997, Boemare 2002), that account for most of the virulence associ-
ated with the nematodes. House fly larvae and adults are highly sus-
ceptible to Steinernema feltiae (=carpocapsae) Filipjev (Rhabditida:
Steinernematidae) and Heterorhabditis heliothidis (=bacteriophora)
(Khan, Brooks and Hirschmann) (Rhabditida: Heterorhabditidae)
under laboratory conditions (Geden et al. 1986).
Applications of entomopathogenic nematodes can target im-
mature or adult house flies. However, successful applications of
nematodes to control immature house flies may depend on larval
development areas. Efficacy in poultry manure is poor (Renn et al.
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
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1985, Georgis et al. 1987, Mullens et al. 1987b, Mullens et al. 1987c,
Archana et al. 2017), but may be better in cattle or swine manure
(Taylor et al. 1998, Renn 1998). Additives, such as calcium alginate
capsules, may allow nematodes to be more effective by reducing un-
intended mortality (Renn 1995). Like entomopathogenic fungi, spe-
cies and strain may influence virulence for fly control (Taylor et al.
1998, Wojciechowska et al. 2013), thus the development of superior
strains and formulations may have greater impact on fly control.
Adult house flies also can be targeted with nematodes (Renn et al.
1999). Delivery method and complementary additives or formu-
lation may be critical to high infectivity (Geden et al. 1986, Renn
1998, Renn and Wright 2000).
Entomopathogenic nematodes are attractive candidates as bio-
logical control agents. They often have a wide host range, can be
easily mass-produced at low cost, have a long storage life (especially
Steinernema spp.), can be selected for desirable traits, are compatible
with certain insecticides, are safe for vertebrates, and are easily ap-
plied in the field. However, efforts will need to be made to conduct
field research to demonstrate utility for house fly control in various
livestock and equine facilities, and improvements to delivery systems
could support their use in poultry facilities.
Black Soldier Fly: a Special Case
Black soldier fly (Hermetia illucens (L.)) is found in temperate and
tropical regions and has received considerable attention for its
ability to decompose animal waste (Gold et al. 2018, Lalander et al.
2019) and convert it to an insect source of animal and human food
(Wang and Shelomi 2017, Beskin et al. 2018, Tomberlin and Van
Huis 2020). Soldier flies are most common in poultry layer facil-
ities, where the activity of their larvae modifies the manure habitat
in ways that make it unsuitable for house fly larvae (Furman et al.
1959, Sheppard 1983). The presence of their larvae in manure also
inhibits oviposition by house flies (Bradley and Sheppard 1984).
Chemical Control
Many chemicals are used in pest management, but differ in
range of action, toxicity, and persistence in the environment.
Insecticides are synthetic or natural compounds that act as direct
toxins to house flies. Control of house flies is most commonly
attempted with conventional insecticides because of their rapid
action, low cost (relative to other strategies) and effectiveness
(at least for new insecticides). However, resistance can radically
reduce the effectiveness of insecticides for fly control. The spe-
cific insecticides available for fly control vary by state (Gerry
2020) (https://www.veterinaryentomology.org/vetpestx). A 2007
survey of U.S. dairies indicated numerous insecticides were used
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1 21

for house fly control in the United States [chlorpyrifos, cyfluthrin,
cyhalothrin, cypermethrin, dichlorvos, diflubenzuron, fenvalerate,
imidacloprid, methomyl, naled, permethrin (some formulations
contain piperonyl butoxide (PBO), pyrethrins + PBO, spinosad,
and tetrachlorvinphos (some formulations contain dichlorvos)],
although there was significant variability in each state. At present,
the main active ingredients used in fly-control insecticides are pyr-
ethroids (e.g., permethrin, β-cyfluthrin, λ-cyhalothrin) in space
sprays/premise treatments and neonicotinoids (imidacloprid,
dinotefuran, and nithiazine) in a variety of baited delivery sys-
tems. Table 4 provides a list of the common insecticides that are
currently used for house fly control.
Insecticide Resistance and Its Consequences
Resistance has developed to all available insecticides used for house
This has caused tremendous concerns at those facilities (one was
cattle and one was poultry) and has growers and extension agents
scrambling for alternatives. Thus, the evolution of resistance is pitted
against the ingenuity of humans to invent new and safe insecticides.
There are few alternative options for house flies. There are, however,
new insecticides that are registered for use on other insects, but not
house flies, and some of these may be effective against house flies.
The Food Quality Protection Act of 1996 assesses risk based on ag-
gregate exposures, and has led to very focused registrations for new
insecticides over the last two decades as companies selected markets
that would offer the greatest profitability. As a result, for the last 20
yr there have been no new insecticides coming to market that were
labeled as premise sprays for house flies in the United States. Thus,
if resistance to pyrethroids renders these insecticides ineffective over
a wide region (beyond the single facility failures that have been

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fly control and is a global problem (Cao et al. 2006, Scott et al. 2013,
Scott 2017). The speed with which a given insect species evolves
resistance can vary considerably, and in some cases is remarkably
rapid (after a single spray season). House flies have consistently been
one of the species in which resistance evolves quickly. Resistance
has evolved rapidly and globally to nearly every class of insecticide,
including organochlorines, organophosphates, carbamates, pyr-
ethroids, insect growth regulators, neonicotinoids and spinosyns
(Keiding 1986, Gerry and Zhang 2009, Scott et al. 2013, Table 4).
Recently, facilities have been identified where complete failure of
house fly control by pyrethroids was observed (Freeman et al. 2019).
observed), there are only older insecticides available to owners of
livestock production facilities as premise treatments. Almost all of
these older insecticides are also plagued by resistance issues (Table 4)
and have unfavorable environmental and mammalian toxicological
profiles (e.g., organophosphates). To complicate matters further,
house flies have tremendous potential for cross-resistance to arise
from overuse of a single chemical (e.g., Wen and Scott 1997, Shen
and Plapp 1990, Abbas et al. 2015, Scott 1989). Baits containing
neonicotinoids and spinosad were highly effective at the time of
introduction but resistance has limited their effectiveness as well
(Table 4).

Table 4. Documented house fly resistance to insecticides with different modes of action

Insecticide or Target Delivery methodsa Cases of resistance in North Americab

insecticide class life stage

Carbamates Adult Spray, bait Forgash and Hansens 1959, Georghiou et al. 1961, Plapp and Bigley 1961,
Georghiou 1962, 1966, Harris et al. 1982, Price and Chapman 1987, Bull
and Pryor 1990, Liu and Yue 2000, Scott et al. 2000, Kaufman et al. 2001b,
Darbro and Mullens 2004, Scott et al. 2013, Freeman et al. 2019
Organophosphates Adult Spray, dust, feed- Fay et al. 1958, Hansens 1958, Labrecque et al. 1958, Forgash and Hansens
through, impreg- 1959, Harris and Burns 1959, Wilson et al. 1959, Forgash and Hansens
nated strip 1960, Hansens 1960, Bigley and Plapp 1961, Labrecque and Wilson 1961,
Georghiou 1962, Georghiou and Bowden 1966, Forgash and Hansens
1967, Georghiou 1967, Mathis et al. 1967, Georghiou and Hawley 1971,
Georghiou et al. 1972, Harris et al. 1982, Boxler and Campbell 1983, Mac-
Donald et al. 1983, Bloomcamp et al. 1987, Bull and Pryor 1990, Liu and
Yue 2000, Scott et al. 2000, Kaufman et al. 2001b, Marcon et al. 2003, Scott
et al. 2013, Freeman et al. 2019
Pyrethrins and Adult Spray, fog/aerosol Forgash and Hansens 1959, Georghiou 1962, DeVries and Georghiou 1980,
Pyrethroids Harris et al. 1982, MacDonald et al. 1983, Scott and Georghiou 1985,
Meyer et al. 1987, Bull and Pryor 1990, Meyer et al. 1990a, Chapman et
al. 1993, Liu and Yue 2000, Scott et al. 2000, Kaufman and Rutz 2001,
Kaufman et al. 2001b, Marcon et al. 2003, Kaufman et al. 2010, Scott et al.
2013, Freeman et al. 2019
Neonicotinoids Adult Bait, spray, impreg- Liu and Yue 2000, Kaufman et al. 2006, Kaufman et al. 2010, Scott et al. 2013
nated strip
Spinosyns Adult Bait, spray Scott et al. 2000, Deacutis et al. 2007
Pyriproxyfen Larvae Spray, fog/aerosol Cerf and Georghiou 1972, Plapp and Vinson 1973, Cerf and Georghiou 1974,
Georghiou et al. 1978, Pimprikar and Georghiou 1979c
Benzoylureas Larvae Spray, feed-through Pimprikar and Georghiou 1979, Shen and Plapp 1990
Cyromazine Larvae Feed-through, spray Pimprikar and Georghiou 1979, Iseki and Georghiou 1986, Scott et al. 2000
Indoxacarb Adult Spray Shono et al. 2004
Diamides Adult Bait None as of April 2020

a
Current and past methods. Some approved methods are rarely used. Sprays = premise, animal and/or manure applications (see https://www.veterinaryentomology.
org/vetpestx for details)
b
Cases of cross-resistance.
c
Cases of methoprene resistance.
22
New Tools for House Fly Management
Background
Control technologies outside of the traditional chemical insecticides
and biological control agents discussed above have long been of
interest to researchers. Household chemicals such as borax, bleach
(chloride of lime), and table salt were studied for addition to manure
to reduce fly production. Early concerns were that such larvicides
might have detrimental effects on manure that might decrease its
value as fertilizer (Cook et al. 1914, Hewitt 1914a,b). Further motiv-
ation for discovering alternative chemical control technologies came
shortly after the DDT product GNB (Gerasol-Neocid-Base) came to
market in 1942. By the late 1940s, studies of DDT failure in house
fly control began to appear in the literature (e.g., Barber and Schmitt
1948, Lindquist and Wilson 1948, Wilson and Gahan 1948). Shortly
after, novel methods to restore the usefulness of some important in-
secticides like DDT and pyrethrum were developed (Wilson 1949).
These came in the form of the insecticidal synergists piperonyl
cyclonene and piperonyl butoxide (PBO), the latter of which is still
widely used in house fly control products. More attention was also
turned to traps and attractants (Yates 1951, Muto and Sugawara
1965).
Current Outlook
Today, with widespread resistance to nearly every insecticide mar-
keted for house fly control (Freeman et al. 2019), discovery of novel
chemistries has never been more important. Chemistries that have
insecticidal activities or synergistic potential with currently used in-
secticides are of particular interest. One broad group of compounds,
the essential oils, have received considerable attention recently
(Pavela and Benelli 2016, Khater and Geden 2019). However, of the
hundreds of studies published on their insecticidal potential per year
for numerous insect pest species, few provide the chemical informa-
tion or positive control comparisons (i.e., comparisons to current
insecticides) necessary for proper evaluation (Isman and Grieneisen
2014). It is therefore imperative that future work include these im-
portant parameters. An active component of eucalyptol, 1,8-cineole,
appears to be among the most potent (Palacios et al. 2009, Rossi and
Palacios 2015). When applied as a fumigant, toxicity of 1,8-cineole
(LC50: 3.3 [1.1–10.4]) mg/dm3, was similar to deltamethrin (LC50:
9.2 [2.8–29.5] mg/dm3). 1,8-cineole also appears to synergize well
with the pyrethroid deltamethrin (deltamethrin + 1,8-cineole: 1.0
[0.08–12.1]) mg/dm3, deltamethrin + PBO: 1.5]0.2–11.4)] mg/dm3).
Other essential oils such as geraniol show some repellent potential
and are sometimes classified as minimum risk compounds (USEPA
2015). Geraniol-based fly repellent sprays are popular on the market
(e.g., Outsmart, Pyranha Zero Bite All Natural).
Few new traditional insecticides have come to market for
house fly control over the past 5 yr. Perhaps the most prominent is
cyantraniliprole, belonging to the second-generation of anthranilic
diamide insecticides (Yu 2015). Insecticides in this class bind to the
insect ryanodine receptor, which is a new mechanism of action that
minimizes the chance for cross-resistance from previous insecticide
use. Cyantraniliprole was registered in a house fly granular bait in
2015 (Zyrox, renamed Cyanarox). In 2018, BASF initiated an EPA
registration of the meta-diamide broflanilide (federal register cit-
ation: 83 FR 34128). Included in the registration is use of broflanilide
in two fly baits (Vedira) targeting multiple filth flies, including house
flies. No new insecticides have been registered as premise treatments
for >20 yr.
Within the past few years, the non-nutritive polyols eryth-
ritol and xylitol have been assessed for their potential as house fly
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
adulticides and larvicides, with mixed results (Burgess and King
2017, Fisher et al. 2017, Burgess and Geden 2019). Under no-choice
conditions, over 50% of flies die within two days when exposed to 2
M solutions of the polyols. However, given acute exposures of 24 h,
followed by access to normal food, fly mortality barely reached 25%
after 20 d, and was not significantly different from the normal food
control. Similarly, larvicidal LC50 values against late second instar
flies were in the parts-per-thousand range for house flies, making
them orders of magnitude worse than typical insect growth regu-
lators. Lower concentrations were very effective at killing eggs/early
instar larvae, with 0% of eggs developing to adults in the erythritol
treatment and only 23.3% of eggs developing to adults in the xylitol
treatment. This is compared to 46.0 % of control eggs developing to
adults. The polyols erythritol and xylitol also appear to be attractive
substitutes for sucrose in baits, especially those centered around the
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entomopathogenic fungus, Beauveria bassiana (Burgess et al. 2018).
Current work suggests that they may also play a role in the house fly
gut physiology and microbiome, potentially leading to new methods
for controlling house flies by exploiting their physiology in ways in-
dependent of traditional insecticides.
House flies are not generally regarded as suitable candidates
for conventional (irradiation-based) sterile insect technique (SIT)
methods. Population densities are too large and fluid to overwhelm
with irradiated males, and male flies are as pestiferous and cap-
able of transmitting pathogens as females. Emerging technologies
may create new opportunities to examine SIT and other genetic
methods (Scott et al. 2018). Promising results with mosquitoes have
been seen using Wolbachia/cytoplasmic incompatibility (Bourtzis
et al. 2014, Mains et al. 2019) and the release of genetically modi-
fied males to deliver dominant lethal alleles (RIDL) (Harris et al.
2012). Moreover, the availability of the genome of house flies (Scott
et al. 2014) and related species may lead to novel management ap-
proaches that exploit vulnerabilities in the life cycle. Little work
has been done using RNAi in adult house flies, but constructs have
been identified that, when injected, shut down ovarian development
(Sanscraint et al. 2018).
Research Priorities
The house fly remains a formidable adversary with many un-
answered questions despite over 100 yr of scientific research. Topics
that warrant highest priority for future research needs fall under
four broad categories.
Management
Monitoring flies remains a highly inexact science. More research
is needed to identify which monitoring tools are most appropriate
in different animal commodity systems. Nuisance/action/economic
thresholds are in urgent need of revisiting, as current threshold
values are based on little more than traditional use that has been
repeated for nearly 50 yr. The use of monitoring house fly density
in the decision making for insecticide applications is sorely needed.
Biological control has won acceptance as a management tool
in some settings, but there are still gaps in availability of products
that are effective and economically feasible. Many mycopesticides
have been developed and labelled for crop pests, and screening
these for house fly control would facilitate label expansion to in-
clude this pest. Dozens of laboratory studies on efficacy of fungal
pathogens for house flies have been conducted, but only a handful
of field tests have been published. Field testing of promising patho-
gens is critically needed to encourage commercial producers to
Journal of Integrated Pest Management, 2021, Vol. 12, No. 1
invest in developing pathogens as viable and competitive products.
One of the main liabilities of fungal pathogens is their slow kill
rate compared to chemical insecticides. Further research is needed
to identify fast-killing strains and improve the virulence of known
strains by selection, genetic modification, and better formulations
either as stand-alone agents or in combination with other compo-
nents and adjuvants. New formulations could also include baits and
attract-and-infect stations that could be used as autodissemination
devices. Other biological control research needs include the develop-
ment of predator mites for augmentative control and improvements
in timing, species selection, and release rate recommendations for
pteromalid parasitoids in different animal systems.
Traps are frequently used by producers (USDA 2014) and will
always play an important role in house fly management, yet there
have been no substantial improvements in house fly attractants for
traps in over 40 yr. Renewed research is needed to identify superior
attractants, especially ones that are not malodorous to humans and
that would preferentially attract female flies.
Novel chemistries are needed to develop new insecticide prod-
ucts to provide rapid reduction of house fly populations resistant
to currently available insecticides. Screening of currently-registered
insecticides not already in use for house fly control is also needed,
with special emphasis on space sprays and modes of action distinct
from those of pyrethroids, neonicotinoids, and organophosphates.
Integrated resistance management (IRM) plans that take an area-wide
approach would be helpful in extending the effective product life of
those insecticides that remain effective. Screening of essential oils and
other plant-derived compounds for insecticidal and repellent potential
should continue, with rigorous testing standards that include testing
against positive controls and inclusion of information on chemical
composition and purity. New insecticide synergists are needed as well,
especially molecules that will fit an OMRI or EPA minimal risk profile.
Rising global temperatures will require fly management to be
conducted under increasingly hot conditions in many locations. High
temperatures can impact the effectiveness of biological (Geden et al.
2019) and chemical control (Scott and Georghiou 1984). Research
on fly management under hot conditions, especially field studies, is
needed to prepare for future challenges.
Biology and Life History
There is no satisfactory answer to the important question of how flies
typically live in the field. Field longevity experiments are notoriously
difficult but are needed to identify factors affecting fly survivorship
under natural conditions. For example, how is longevity affected by
animal production systems, food resources within those systems, and
locally available microbiomes? Research on fly functional genomics,
transcriptomics, and proteomics, leveraged by knowledge gained in
other dipteran pests, may reveal as-yet undiscovered fly vulnerabil-
ities that can be exploited. We are just beginning to explore the roles
that microbiomes play in larval and adult fly fitness. Further research
on this topic is critically needed and could point to ways to interfere
with the acquisition and retention of needed microbes. Several im-
portant aspects of fly behavior are still poorly understood, especially
regarding flight activity and dispersal of flies. A better understanding
of the role of fly sex, local availability of food resources, and fly re-
sponse to environmental conditions could help identify triggers that
lead to dispersal and increased synanthropy.
Role as Reservoirs and Transmitters of Pathogens
Flies that originate from concentrated animal feeding and produc-
tion operations can disseminate microbes within the operations
23
(creating an animal health concern) or away from operations (cre-
ating a human health concern). The studies listed in Table 3, as well
as many other surveys (reviewed in Nayduch and Burrus 2017),
demonstrate that wild-caught flies harbor a diverse and abundant
community of human and animal pathogens in a variety of settings.
Furthermore, the studies we review above demonstrate that flies
can acquire, harbor, and transmit a number of pathogenic bacteria,
and may facilitate lateral transfer of resistance and virulence genes
among microbes in the gut. Several critical research gaps remain
in defining the role house flies play as both reservoirs and trans-
mitters of microbes. What are the most important microbial and
fly-specific variables that impact acquisition, persistence and trans-
mission of pathogens? What factors underlie the temporal dynamics
of pathogen persistence in infected flies? How long do flies harbor
pathogens? How long do they shed pathogens? Are pathogens being
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shed in doses sufficient to infect animals or humans? How signifi-
cant a role does house fly play in contaminating human foods/crops
with pathogens? A better understanding of these factors, along with
variables from prior studies outlined in this review, should be used
to develop predictive models as improved risk assessment tools for
transmission of animal and human pathogens. Further demonstra-
tions also are needed to document that effective fly management
mitigates pathogen prevalence in the field.
Systems Needs
Intensive animal agriculture production systems provide materials
that attract adult house flies and allow for the development of im-
mature house fly populations on site. These include animal ma-
nures, collected and stored, and feed ingredients, many of which
are suitable for house fly development. These materials are continu-
ally replenished. Fly management programs in all intensive system
focus on fly habitat management/reduction, and the available man-
agement tools are applicable across different animal production
systems. However, additional research is needed to develop more
effective manure and feed storage systems that limit fly access and
are easy to use.
Equine systems have received the least research attention with re-
gard to house fly management. Fly populations tend to be compara-
tively low, and many of the flies present may have been produced
off-site on cattle or other facilities. Research is needed to determine
high-risk larval development habitats and to assess the proportion
of flies on horse farms that have developed on-site. Many equine
facilities use fly parasitoids, yet almost nothing is known about the
efficacy of parasitoid releases on horse farms. There are currently no
scientifically based recommendations for the numbers or even the
species of parasitoids needed to keep flies at acceptable levels in this
difficult market.
The poultry industry is changing rapidly and is now composed
of organic and conventional production systems using a wide array
of housing types, flock sizes, and management practices. Much
of the research on house fly management has been conducted on
farms with high-density caged layer houses that house >100,000
hens each. Europe and California have banned this type of housing,
and future fly control work will be conducted on smaller open-floor
facilities of different designs. Research is needed to assess how fly
populations are affected by housing type and husbandry practices
in this rapidly evolving industry. The swine industry faces similar
challenges, with a range in production types and over wide geo-
graphic areas. Best management practices are critically needed in
the poultry and swine industries for fly monitoring and biological
control options under different housing, management, and climate
conditions.
24
Acknowledgments
We thank members of the S1076 multistate project ‘Fly Management in Ani-
mal Agriculture Systems and Impacts on Animal Health and Food Safety for
manuscript discussion and advice. We also appreciate the financial support
from Penn State Extension grant to hold a workshop in Orlando, Florida,
for the discussion of this manuscript and others in the series. We thank Dana
Johnson and Brianna Davis for help in organizing the references and the fol-
lowing individuals for reading and editing the manuscript before it was sub-
mitted: Fallon Fowler, Bethia King, David Taylor, Jerome Hogsette, Roger
Moon, and Kateryn Rochon. Mention of trade names or commercial products
in this publication is solely for the purpose of providing specific information
and does not imply recommendation or endorsement by the U.S. Department
of Agriculture. USDA is an equal opportunity provider and employer.
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