Journal of Economic Entomology, 113(6), 2020, 2571–2580

doi: 10.1093/jee/toaa229
Advance Access Publication Date: 15 October 2020
Review Review

Review of Methods to Monitor House Fly (Musca

domestica) Abundance and Activity
Alec C. Gerry

Downloaded from https://academic.oup.com/jee/article/113/6/2571/5924105 by Universitas Indonesia user on 27 October 2023

Department of Entomology, University of California, 900 University Avenue, Riverside, CA 92521 (alec.gerry@ucr.edu)

Subject Editor: Jeb Owen

Received 6 July 2020; Editorial decision 29 August 2020

Abstract
The house fly is a ubiquitous pest commonly associated with animal facilities and urban waste. When present
in large numbers, house flies can negatively impact humans and animals through nuisance and the transmis-
sion of pathogens. Since the development of fly traps and sticky papers to capture flies in the late 1800s, these
and other methods have been used as a means to monitor change in house fly density or fly activity over time.
Methods include substrate sampling to record density of immature flies, visual observations of adult fly ac-
tivity, instantaneous counts of landing or resting flies, accumulation of adult flies on/in traps, or accumulation
of fly fecal and regurgitation spots deposited by flies onto white cards. These methods do not estimate true
house fly density, but rather provide an index of house fly activity that is related to both fly density and the fre-
quency of individual fly behavior (e.g., frequency of flight, landing events) and which is likely more predictive
of negative impacts such as nuisance and pathogen transmission. Routine monitoring of house fly activity is
a critical component of a house fly management program. Fly activity should be held to a level below a pre-
determined activity threshold ('action threshold') above which negative impacts are anticipated to occur. This
article is a review of methods utilized for monitoring house fly (Diptera: Muscidae) activity.

Key words: surveillance, trap, density, threshold, precision

The house fly (Musca domestica L.) is a ubiquitous pest commonly
associated with animal facilities where this fly develops in the feces,
wet feed, and other decaying organic matter that is often abundant
at these facilities (Geden and Hogsette 1994). House flies are also as-
sociated with urban sites including waste handling facilities (Howard
2001, Lole 2005). When produced in large numbers, house flies can
cause considerable nuisance to surrounding communities poten-
tially resulting in costly citations, fines, and lawsuits (Thomas and
Skoda 1993, Winpisinger et al. 2005). The house fly is also an im-
portant mechanical vector of human and animal pathogens (reviews
by Howard 1911, West 1951, Wolf and Zijl 1969, Greenberg 1971,
Olsen 1998, Nayduch and Burrus 2017). Incidence of human disease
can be reduced following implementation of area-wide house fly
management programs (Watt and Lindsay 1948, Lindsay et al. 1953,
Lindsay and Scudder 1956, Cohen et al. 1991, Levine and Levine
1991, Chavasse et al. 1999, and reviewed by Greenberg 1973).
Animal production and urban waste handling facilities should
strive to maintain adult house fly activity below an ‘action threshold’
above which economic injury or negative impacts (i.e., nuisance or
pathogen transmission) might occur (Stern et al. 1959, Flint and van
den Bosch 1981). Ideally, fly activity is routinely monitored so that
control measures are applied before fly activity exceeds the predeter-
mined action threshold (Urech 2004). Monitoring of fly activity may
be required by federal or local health agencies. For example, moni-
toring of fly activity is mandated for egg-layer facilities as part of
the U.S. Food and Drug Administration Salmonella Enteritidis pre-
vention strategy (FDA 2009). Unfortunately, there are no accepted
standards for methods to monitor house fly activity and suitable
action thresholds are even more elusive. Nevertheless, routine moni-
toring of house fly activity can provide an important record to ad-
dress fly nuisance complaints or lawsuits against a facility.
The lack of accepted standards for monitoring house fly ac-
tivity presents challenges to facilities that desire (or are required)
to monitor fly activity, thus discouraging implementation of an ef-
fective fly monitoring program. In the absence of a fly monitoring
program, fly control is often initiated only when adult fly activity
has resulted in obvious negative impact to the facility or to neigh-
bors, suggesting that an action threshold of adult fly activity is al-
ready exceeded and application of insecticides targeting adult flies
is required (Axtell 1970a, Geden and Hogsette 1994). Also without
a fly monitoring program, measures applied to control flies cannot
be evaluated for effectiveness, other than through qualitative assess-
ments of fly activity (e.g., fewer reports of nuisance). This can lead to
continued use of ineffective measures such as insecticides to which
house flies have developed a high degree of resistance (reviewed by
Keiding 1999, Freeman et al. 2019). The lack of standards for house

© The Author(s) 2020. Published by Oxford University Press on behalf of Entomological Society of America. 2571
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2572
fly monitoring methods also impacts development of new fly con-
trol products and novel insecticides since efficacy evaluations vary
among studies making comparisons difficult.
This article is a review of methods utilized for monitoring house
fly activity and includes discussion of the application and challenges
to their use primarily in animal production systems.
House Fly Monitoring Methods
Methods for monitoring house fly activity have been described for
poultry houses (Anderson and Poorbaugh 1964; Axtell 1970a,b;
Burg and Axtell 1984; Beck and Turner 1985; Lysyk and Axtell
1986), cattle feedlots (Urech et al. 2004), dairies (Pickens et al.
1972, Pickens and Miller 1987, Gerry et al. 2011), swine housing
(Machtinger and Burgess 2020), and horse facilities (Machtinger
et al. 2016). Monitoring house fly activity can focus on any life stage
of the house fly, but activity of adult flies is usually of greatest interest
since nuisance and pathogen transmission are negative impacts as-
sociated with the adult life stage. Monitoring methods discussed in
this article do not estimate the true size of the house fly population,
as this would require more complex methods (Southwood 1978,
Kristiansen and Skovmand 1985), but rather these methods provide
a measure or index of house fly activity that is related to both fly
density and fly behavior (e.g., frequency of flight, landing habits, re-
sponse to odors or visual cues). Since the negative impacts of flies,
including nuisance and pathogen transmission, are also likely related
to both fly density and the frequency of fly behaviors, an index of
fly activity is likely to be more predicative of negative impacts than
would be an estimate of fly density alone.
Immature Sampling
Sampling of accumulated animal feces or other developmental sub-
strate for the presence of immature flies may give an early indica-
tion of future adult fly abundance and activity. A garden trowel or
soil coring tool can be used to acquire substrate samples of similar
volume and depth (usually the top few inches of substrate) which
are then spread out within a pan to observe and count immature
flies by species (Brydon 1966, Eastwood et al. 1967, Smith and Rutz
1991, Hogsette et al. 2012). Immature flies can also be separated
from the substrate sample using a Berlese funnel (Brydon and Fuller
1966, Axtell 1970a, Stafford and Bay 1987, Schmidtmann 1988),
by washing substrate through sieves to retain immatures (Eastwood
and Schoenburg 1966, Schoenburg and Little 1966), or by submer-
ging substrate in water containing high-salt concentration (e.g.,
using magnesium sulfate) (Ladell 1936, Southwood 1978) causing
immature flies to float to the water surface where they are readily
collected (Laurence 1954, Matthysse and McClain 1973, Pitts et al.
1998). Water with a high-sucrose concentration can also be used
to separate immature flies from a substrate sample, at least for re-
covery of third-instar house fly larvae (Tobin and Pitts 1999). The
salt flotation method is reported to be less efficient for collection
of immatures compared with washing substrate through sieves
(Eastwood and Schoenburg 1966). One potential benefit of the salt
flotation method is that separated immatures remain alive and can
be placed into suitable media to be reared to the adult stage to con-
firm species identification (Ladell 1936, Laurence 1954). Eggs and
early instar fly larvae can be difficult to see in substrate samples
and are often excluded from substrate sample counts (e.g., Stafford
and Bay 1987, Tobin and Pitts 1999). Since the cumulative mor-
tality of eggs and early instar larvae can be high (Morgan et al.
1981), the abundance of late-stage house fly larva and pupae in
Journal of Economic Entomology, 2020, Vol. 113, No. 6
substrate samples is perhaps a better predictor of future adult fly
activity anyway.
A method similar to immature sampling is the capture of recently
eclosed adult flies as they emerge from a substrate after completing
their immature development. Emerging adult flies are captured
using an emergence trap placed onto the substrate that directs the
emerging adult flies by way of a screen or funnel into a collection jar
(Wright et al. 1973). Emergence traps have also been used as a com-
panion tool for immature sampling, with substrate samples placed
into containers fitted with an emergence trap to collect emerging
adult flies (Meyer et al. 1987, Mullens et al. 1996, Cook et al. 1999).
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Immature sampling is limited as a method for routine monitoring
of house fly activity given the variance in immature density among
small substrate samples (Schoenburg and Little 1966; Stafford and
Bay 1987, 1994) and the need to estimate immature mortality if the
aim is to predict future adult fly activity. While emergence traps may
improve prediction of future adult house fly activity since an esti-
mate of immature mortality is not needed, this has not been tested.
Nevertheless, immature sampling methods remain useful for other
purposes, such as to identify productive house fly developmental
sites for treatment or to evaluate fly production potential of different
substrates or substrate conditions (Eastwood et al. 1967, Axtell
1986, Stafford and Bay 1987, Schmidtmann 1988, Gerry et al. 2005,
Hogsette et al. 2012).
Scudder Fly Grid
This device, developed by Harvey Scudder of the U.S. Public Health
Service, is reported to be a ‘standard neutral resting surface’ for adult
house flies (Scudder 1947). The grid is constructed of wooden strips
fixed to a 3.0 ft2 (0.91 m2) frame with open slits between the wooden
strips providing edges attractive to resting house flies while also not
obscuring the attractive substrate upon which the monitoring device
is placed. The grid is placed on the ground at a location where fly
activity is noticeably concentrated, and then flies resting on the sur-
face of the grid are counted 30 s later. The fly grid therefore provides
an instantaneous record of fly activity at the location and time that
the observation is performed. Because the fly grid is placed in areas
of concentrated fly activity, the count is an index of fly activity not
a measure of fly density (Murvosh and Thaggard 1966). Scudder fly
grid counts were highly related to visual counts of flies recorded at
the same location immediately before the grid was placed (Welch
and Schoof 1953), but the grid ensures a standard visual target and
is therefore expected to provide greater consistency in fly counts
among sites and observers.
Scudder used the highest of five consecutive grid counts as
the fly activity count of record for a single location and time, but
Schoof (1955) suggests that only one grid count per attractive site
is needed since additional counts will usually have lower fly num-
bers due to repeated disturbance of the flies (see also Holway et al.
1951). Murvosh and Thaggard (1966) reported that waiting only
10 s before counting flies on the grid will give a higher fly count since
landing flies quickly crawl down between the wooden strips to reach
the attractive substrate beneath the grid. Dhillon and Challet (1985)
support using a shorter time from grid placement until recording the
fly count, finding that 50% of flies land on a grid within the first 15 s
during a 60-s observation period.
The Scudder fly grid is reported to be a satisfactory index of
adult house fly activity (Watt and Lindsay 1948, Murvosh and
Thaggard 1966) and the fly grid continues to be used to monitor
fly activity (Morris and Hansen 1966; Morgan et al. 1975,
1981; Morgan 1980; Krafsur 1985; Crespo et al. 1998), with
Journal of Economic Entomology, 2020, Vol. 113, No. 6
methods used generally following those outlined by Murvosh and
Thaggard (1966). However, in several studies where Scudder grid
counts were compared with other measures of fly activity, the grid
counts were not related to fly density or to other measures of fly
activity (Logan 1953, Pickens et al. 1972, Beck and Turner 1985,
Lole 2005), suggesting caution when considering this method for
routine monitoring of adult house fly activity. House fly activity
varies by time of day and with environmental conditions (Parker
1962, Zahn and Gerry 2020), and an instantaneous record of
fly activity such as a grid count is expected to reflect conditions
at the time of sampling (Lysyk and Moon 1994). Additionally,
using the Scudder fly grid where several fly species are abundant
can be a challenge, given the difficulty to identify at a glance the
flies landing on the device.
Sticky Fly Ribbons
The use of ribbons, tapes, and papers coated with a sticky sub-
stance to capture resting flies has a long history. A patent for sticky
fly paper was first recorded in 1908 (Ross 1908) and Howard
(1911) describes using suspended sticky papers to take advan-
tage of house fly behavior to preferentially land on hanging ob-
jects such as curtain cords and suspended lamp chains. The coiled
sticky fly ribbon still in use today was patented shortly thereafter
(Wagner 1919). House flies prefer to land on longer, thinner fly
papers (750 × 20 mm) relative to shorter, wider fly papers (375 ×
40 mm) (Raybould 1964) perhaps supporting the observation that
house flies are attracted to surface edges (Scudder 1947). Fly rib-
bons will capture several pest fly species, allowing activity of these
species to be monitored simultaneously (Anderson and Poorbaugh
1964; Axtell 1970a,b; Legner et al. 1973; Lysyk and Axtell 1986;
Gerry et al. 2011).
Fly ribbons are best placed at locations frequented by flies
(Pickens et al. 1972), such as indoors near the roof of animal housing
where flies accumulate at night (Anderson and Poorbaugh 1964).
Most researchers report good fly capture when fly ribbons are placed
to hang from midline roof supports within poultry houses or cattle
barns (Anderson and Poorbaugh 1964; Axtell 1970a,b; Pickens et al.
1972; Legner et al. 1973; Rutz and Axtell 1979; Quisenberry and
Foster 1984; Lysyk and Axtell 1986; James et al. 2017). Dusty condi-
tions reduce the stickiness of fly ribbons making them unsatisfactory
for monitoring fly activity in broiler-breeder poultry facilities (Rutz
and Axtell 1981) or drylot dairies (Gerry et al. 2011). Fly ribbons ex-
posed to direct sun will also quickly lose their stickiness (Anderson
and Poorbaugh 1965).
Fly activity counts using fly ribbons hung from roof rafters in
open-sided poultry houses for 24 h were positively correlated with
the number of flies emerging from poultry manure collected from
these same poultry houses, suggesting that fly ribbons are a suitable
means to measure fly density (Legner et al. 1973). Fly ribbons hung
for 24 h provided a more reliable index of changing house fly density
than were fly ribbons hung for only 1–2 h (Pickens et al. 1972, Beck
and Turner 1985), and 24-h fly ribbon counts successfully reflected
a doubling of fly density, whereas 2-h fly ribbon counts, Scudder fly
grid counts, and visual observation of flies resting at a specific pre-
determined location (‘station counts’) failed to detect doubling of fly
density (Pickens et al. 1972). Short period (1–2 h) sampling methods
are very likely subject to the same effects of environmental condi-
tions as described above for the Scudder fly grid. When fly ribbon
sampling periods were increased to ≥2 d, fly counts were related to
other measures of adult fly activity (Rutz and Aztell 1979, Lysyk
and Axtell 1986) as flies accumulated on the fly ribbons over the
2573
full range of environmental conditions experienced during the sam-
pling period (Lysyk and Moon 1994). Fly ribbons have been used to
monitor house fly activity for up to 7 d (Axtell 1970a,b; Lysyk and
Axtell 1986; Turner and Ruszler 1989), but fly capture rates can
be reduced as sticky ribbons fill with flies (Axtell 1970a) suggesting
shorter sampling periods may be needed in facilities with high fly
activity.
Fly ribbons have also be used to acquire a near-instantaneous
fly count using the moving sticky tape or walking sticky tape count
method (Turner and Ruszler 1989). This method was developed for
rapid assessment of fly activity at caged-layer poultry facilities where
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birds are held in rows of suspended wire cages and bird feces ac-
cumulates beneath the rows of caged birds. To perform a moving
sticky tape count, a fly ribbon is held by hand with the bottom of
the ribbon suspended approx. 2.5–5 cm above the ground while the
operator walks at a normal pace between rows of caged birds. Flies
disturbed into flight by the passing movement may be captured on
the fly ribbon. Turner and Ruszler (1989) reported walking a route
to sample an entire poultry house or at least two full cage rows
within the house for a minimum total walking distance of 304.8 m
(1,000 ft). In the study by Turner and Ruszler (1989), fly activity esti-
mated by the moving sticky tape method was not correlated with fly
activity estimated by 7-d fly ribbon counts. The authors suggest that
the moving sticky tape method is an improvement over placing fly
ribbons in fixed locations because the moving tape method captures
flies along the entire walking route, thereby addressing variation in
fly density throughout the poultry house. However, like other in-
stantaneous fly counts, the moving sticky tape method would cer-
tainly be subject to diel and environmental variation in fly activity
perhaps resulting in the lack of correlation between the two sam-
pling methods.
Sticky Fly Traps
Like the fly ribbons described above, sticky fly traps are coated with
a sticky material to capture flies landing on the trap or encountering
the trap during flight. Relative to fly ribbons, sticky traps often have
a larger surface area and many traps have a rigid plastic or fiber-
glass frame. One of the more commonly used sticky traps for moni-
toring fly activity at animal facilities is the Alsynite biting fly trap
(Williams 1973, Broce 1988). Although the Alsynite biting fly trap is
commercially marketed to capture stable flies (Stomoxys calcitrans
(L.)) associated with cattle or horse facilities, it has also been used
to monitor house fly activity at cattle facilities (Geden 2005, Gerry
et al. 2011, Urech et al. 2011), horse facilities (Machtinger et al.
2016), and in urban environments (Winpisinger et al. 2005). Other
sticky fly traps that have been used for monitoring fly activity include
sticky cards (Black and Krafsur 1985, Hogsette et al. 1993, Geden
et al. 1999), sticky cans (Kaya and Moon 1978, Black and Krafsur
1985), sticky pyramids (Pickens and Miller 1987), and sticky panels
(Kaufman et al. 2001, 2005; Zahn and Gerry 2020). While any of
the numerous commercially available sticky fly traps might similarly
be used for fly monitoring, these traps are generally untested for effi-
cacy as fly capturing or monitoring devices.
The placement and use of sticky traps will vary with the many
trap designs in this category of fly monitoring method. As for fly
ribbons, it is expected that longer sampling periods will provide
a better index of overall fly activity as compared with sampling
periods ≤24 h. Relative to fly ribbons, many sticky traps may be
more suitable for outdoor use and can often be deployed for longer
sampling periods given their rigid frame and larger surface area. But
like fly ribbons, sticky traps may be ineffective in dusty conditions
2574
(e.g., Kaufman et al. 2001, Gerry et al. 2011). Also, removal of cap-
tured flies by wild birds can impact activity estimates (e.g., Gerry
et al. 2011).
Attractant-Baited Traps
A patent filed in 1872 described a fly trap to be placed above food
waste, offal, or other fly-attractive materials to capture flies by
directing them upward through an inverted wire mesh cone into a
larger screened cage (Harper 1872). Though versions of this conical
fly trap existed prior to 1872, Harper’s patent was well timed as
scientists were just beginning to appreciate flies as carriers of patho-
gens (e.g., Howard 1911). In the early 1900s, Harper’s inverted cone
trap, and many homemade versions, became widely used, and public
health agencies and local businesses even sponsored contests for
community residents to construct and deploy these traps to capture
and kill disease-transmitting flies (e.g., Daubs 2015). While these
traps proved effective at capturing large numbers of adult flies, they
were not particularly efficient for monitoring adult house fly activity
as traps were often baited with food waste or other materials at-
tractive to many pest fly species. Nevertheless, entomologists suc-
cessfully used these attractant-baited traps to capture flies in early
house fly dispersal studies (e.g., Parker 1916; Bishopp and Laake
1919, 1921; Quarterman et al. 1954a,b; Schoof et al. 1952; Schoof
and Siverly 1954a,b) and a USDA issued Farmer’s Bulletin in 1916,
revised in 1921, described several designs for attractant-baited traps
and bait materials that could be used to attract flies (Bishopp 1921).
At present, there are numerous commercially available trap de-
signs that rely on noxious odors to attract and capture flies, with
many traps drowning the captured flies in a smelly liquid solution
within the trap. These ‘stinky fly traps’ can be quite effective for cap-
turing large numbers of flies, but identifying the drowned and rot-
ting flies is difficult and unpleasant. Placing a separate collecting bag
or mesh insert into these traps to separate captured flies from the
liquid solution may simplify use of these traps for monitoring fly
activity (Urech et al. 2011, James et al. 2017). However, the effect-
iveness of attractant-baited traps can vary considerably by proximity
to natural attractants or fly development sites (Pickens et al. 1967,
Pickens and Miller 1987) and baited traps can vary in attractive-
ness by trap design, bait formulation, bait age, and environmental
conditions (Madwar and Zahar 1951, Geden 2005, Geden et al.
2009) potentially making these traps inconsistent for routine house
fly monitoring. Fly age may also be a factor in attraction to baited
traps as suggested by Gerry et al. (2011). Variability in attraction of
baited traps is perhaps responsible for the lack of agreement in fly
activity recorded for baited traps and other trap methods in some
studies (e.g., Schoof 1955). A more uniform attractant such as vin-
egar (Qian et al. 2013), molasses (Bishopp 1921, Brown et al. 1961,
Geden 2005, Geden et al. 2009), or fly-attractive volatiles (Frishman
and Matthysse 1966, Mulla et al. 1977, Hwang et al. 1978, Cossé
and Baker 1996, Quinn et al. 2007, Hung et al. 2019) may reduce
variability among individual baited traps and improve their use for
monitoring fly activity.
To reduce variability among traps and to simplify identification
and enumeration of captured flies, traps can be baited with com-
mercially produced insecticidal fly bait. Captured flies feed on the
insecticidal bait and die within the trap without the need for a liquid
solution to drown the flies. For example, a translucent plastic 1 G
milk jug containing 25 g of dry insecticidal granular fly bait (‘baited
jug trap’) has been used to monitor house fly activity in poultry
houses (Rutz and Axtell 1979, 1981; Burg and Axtell 1984). These
traps are more specific for capture of house flies relative to traps
Journal of Economic Entomology, 2020, Vol. 113, No. 6
baited with waste food or noxious odors (Burg and Axtell 1984).
Baited jug traps are typically placed near the roof peak in narrow
poultry houses, at the level of the highest poultry cages in multilevel
poultry houses, or about 1 m above accumulated feces in high-rise
(deep-pit) poultry houses (Rutz and Axtell 1979, 1981; Burg and
Axtell 1984; Lysyk and Axtell 1985, Stafford et al. 1988). Trap
counts vary along the length of a poultry house (Burg and Axtell
1984) and attractant-baited traps placed at ground level near the
end of a cage row outperformed traps near the middle of the cage
row (Wilson and Mulla 1975), suggesting that trap position even
within a single animal housing structure is important to consider
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when using baited traps. Trap color can also impact fly capture, so
consistency of color is important (Burg and Axtell 1984). In loca-
tions where dust accumulation limits the use of sticky ribbons and
sticky traps, baited jug traps may be the best choice for monitoring
fly activity (Rutz and Axtell 1981).
Baited jug traps hung from the roof of poultry houses for 24 h
provided activity estimates related to fly density (Lysyk and Axtell
1985). And fly counts from baited jug traps hung from the roof for 2
d were related to fly counts using sticky fly ribbons hung for 2 or 7
d (Rutz and Axtell 1979, Lysyk and Axtell 1986). In poultry houses,
baited jug traps were a more reliable indicator of fly density than
were instantaneous counts of resting flies on walls and structural
posts (Stafford et al. 1988). Other traps containing insecticidal fly
bait have been similarly used to monitor fly activity, including place-
ment of an insecticidal bait strip above a fly collection pan (Geden
2005) or addition of dry granular bait to a 5 G (18.9 liter) bucket
(Gerry et al. 2011). Essentially, any collecting device containing in-
secticidal fly bait can be used as long as flies feed on the bait and
the insecticide used produces rapid mortality (Zahn et al. 2019), so
that flies cannot escape the trap. However, use of traps containing
insecticidal fly bait for long-term fly monitoring programs should be
considered carefully since selection for insecticide resistance can be
rapid in house flies (Keiding 1999, Kaufman et al. 2010, Hubbard
and Gerry 2020) and any increase in fly resistance to the insecticidal
bait will confound estimates of fly activity over time.
Spot Cards
Spot cards (or fly speck cards) are white index cards placed at loca-
tions where flies are noted to commonly deposit fecal and regurgi-
tation spots (‘fly spots’), with the number of spots deposited onto
the card over the sampling period recorded as an index of fly ac-
tivity (Axtell 1970a,1986). Any size white index card will work as
a spot card, though 3 × 5 in (7.62 × 12.7 cm) cards are most com-
monly used. If cards of different sizes are used, fly spot counts can
be transformed to spots/cm2 of card area for comparative analysis
(e.g., Gerry et al. 2011). White strips of paper, white tiles, or other
white objects on which flies will similarly land and deposit spots
can also be used (Matthysse and McClain 1973). Several fly species
can deposit fly spots onto these cards, so the spot count is a relative
measure of overall fly activity (Axtell 1970a, Pickens et al. 1972,
Lysyk and Axtell 1986). While the relative contribution of any one
fly species to the total spot card count can perhaps be estimated by
the relative density of each fly species on fly ribbons or using other
adult fly sampling methods (Lysyk and Axtell 1986), this has not
been tested and spot deposition rates may vary among fly species.
While fly ribbons are typically hung from roof rafters, placing
large spot cards to hang vertically from roof rafters for 24 h proved
unreliable as a measure of fly density in dairy barns (Pickens et al.
1972) and poultry houses (Beck and Turner 1985). Spot cards
placed flush against roof rafters for 24 h were even less sensitive
Journal of Economic Entomology, 2020, Vol. 113, No. 6
to changing fly density, particularly as daily temperature increased
and flies increasingly avoided the roof area (Lysyk and Axtell 1985).
Spot cards placed on cage row feed troughs in poultry houses for
24 h provided counts that were more sensitive to fly density even
if spot counts were lower (Lysyk and Axtell 1985). In ventilated
animal housing, spot cards placed on the downwind side of building
support posts provided higher spot counts relative to cards placed
on the upwind side of support posts (Geden et al. 1999).
Increasing the spot card sampling period to 7 d generally im-
proves the relationship of spot card counts to other measures of fly
activity. In poultry houses, spot cards hung from roof supports for
7 d gave spot counts correlated with fly counts from fly ribbons and
baited jug traps hung from roof supports for 2–7 d (Rutz and Axtell
1979, Lysyk and Axtell 1986), and spot cards placed on support
posts for 7 d gave spot counts correlated with fly captures on sticky
cards placed on the posts for just 24 h (Geden et al. 1999). At swine
facilities, counts from spot cards were correlated with counts from
sticky cards when both were placed ~2 m above swine pens over
the same 7-d sampling period (Machtinger and Burgess 2020). At
drylot dairies, 7 d spot card counts were correlated with same week
fly ribbon counts and 1–2 wk later baited trap counts (Gerry et al.
2011). The lag in the fly activity estimate using baited traps relative
to spot cards perhaps suggests that baited traps attract older flies
relative to the presumably age-neutral spot cards, but this should
be more carefully examined since this was noted in only two of the
three dairies sampled.
Perhaps, the biggest challenge to using spot cards as a routine fly
monitoring tool is the time that must be committed to counting the
hundreds or even thousands of fly spots that can accumulate on each
card during the sampling period. To reduce this time commitment,
a spot counting algorithm (FlySpotter) has been developed to iden-
tify and count fly spots on the scanned image of a spot card (Gerry
et al. 2011). The FlySpotter program is available here: https://www.
veterinaryentomology.org/flyspotter-house-fly-monitoring).
Action Thresholds
In crop pest management, negative impacts that result from pest ac-
tivity are largely economic (e.g., loss in profit) and can be readily
modeled against costs to control the pest. This allows for determin-
ation of a pest density action threshold where it is economically ad-
vantageous to control the pest (Stern et al. 1959). For house flies,
negative impacts are largely nuisance, though other impacts such as
spotting of walls, fouling of foods, or transmission of pathogens may
occur in some situations (Axtell and Arends 1990). While high fly ac-
tivity can certainly result in economic costs (e.g., citations, lawsuits,
health expenses), these costs are largely unpredictable. In addition,
many impacts are not strictly economic (nuisance, public relations,
disease). Given the unpredictable nature of economic costs and the
many noneconomic impacts, the development of an appropriate ac-
tion threshold for house fly activity has proven difficult. Given the
association of house flies with nuisance, the term ‘nuisance threshold’
is occasionally used to indicate the fly activity level above which
nuisance complaints may occur (e.g., Quisenberry and Foster 1984).
Even if house fly activity could be modeled against costs, there re-
mains an intractable barrier to determining an action threshold. Fly
nuisance is arguably the most common negative impact of flies, yet
the tolerance for flies varies for each person according to their ex-
perience and expectations. For this reason, many public health agen-
cies simply rely on public complaints of flies as evidence of nuisance
(Thomas and Skoda 1993). However, this approach is unsatisfactory
2575
since it fails to provide a target level of fly activity that must be
achieved to avoid further complaints or at least to avoid a future
determination of nuisance by the local health agency.
To support a more criterion-based fly management program, ac-
tion thresholds are needed to guide control efforts. To address this
need, some researchers have suggested action thresholds that were
subjectively derived from fly activity surveys interpreted through the
professional experience of the researcher. Axtell (1970a) offered the
first fly activity action thresholds for caged-layer poultry farms in
North Carolina. He suggested that fly activity was acceptable below
200 flies/ribbon/week or 100 spots/card/week. These action thresh-
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olds combined counts for several pest fly species since house flies and
little house flies (Fannia canicularis (L.)) were abundant on many of
the farms surveyed. Following a subsequent survey, Axtell (1970b)
estimated appropriate action thresholds to be 100 flies/ribbon/week
or 100 spots/card/week, demonstrating that he understood these ac-
tion thresholds were not universal but were instead specific to the
facilities surveyed, the monitoring methods used, and specific place-
ment of monitoring devices at each facility (Lysyk and Axtell 1986).
Other subjectively derived action thresholds have been suggested as
300 flies/ribbon/week (Lysyk and Axtell 1986), 50 spots/card/week
(Axtell 1986, Lysyk and Moon 1994), and either 350 flies/trap/week
(Rutz 1981, Axtell 1986) or 300 flies/trap/week (Lysyk and Moon
1994) for baited jug traps.
Optimum Sample Size and Sampling Precision
The action threshold can also guide how many sampling devices of
any selected type should be deployed at a facility (‘optimum sample
size’). At fly activity levels near the action threshold, sampling de-
vices must provide enough sampling precision to distinguish a suf-
ficient change in fly activity. Sampling precision is a function of the
mean and variance of sample data, and is related to the density and
distribution of flies and the design and number of sampling devices
used (Southwood 1978). Precision is given by the coefficient of vari-
ability (CV) = SE / x̄, with SE = standard error of the mean, and
x̄ = sample mean. A CV ≤ 0.25 is sufficient to detect a doubling of
fly activity (Southwood 1978), though a CV ≤ 0.15 has been sug-
gested for use given the greater sampling precision (Lysyk and Axtell
1986). Following the method of Karandinos (1976), the expression
above can be rearranged to give n = s2 / x̄ 2(CV)2, where n = sample
size, s2 = sample variance, and x̄ = sample mean. For any monitoring
method, a preliminary survey of fly activity performed on the same
or perhaps similar facility using the desired monitoring method will
provide data to model the relationship of sample mean to sample
variance. With a strong relationship of mean to variance for the
sample data, variance can be expressed as a function of the mean to
replace variance (s2) in the equation above. The minimum number of
sampling devices needed to detect a doubling of fly activity near the
action threshold is then determined by solving for n using CV = 0.25
(or CV = 0.15 if greater precision is desired) and setting the sample
mean ( x̄) to equal the action threshold. Lysyk and Axtell (1986) pro-
vide an excellent review of this method.
Following the methods described above using a CV = 0.15 at
action thresholds of 200 flies/ribbon, 100 spots/card, or 350 flies/
baited jug trap, Lysyk and Axtell (1986) determined that a fly ac-
tivity monitoring program in a narrow caged-layer poultry house re-
quires a minimum of 6 fly ribbons, 7 spot cards, or 9 baited jug traps
(but only 4 baited jug traps in Lysyk and Axtell 1985), whereas a
high-rise (deep-pit) poultry house would require 14 fly ribbons, 8 spot
cards, or 10 baited jug traps. On drylot dairies, Gerry et al. (2011)
2576
determined that 16 fly ribbons, 12 spot cards, or 7 baited traps are
needed to achieve a CV = 0.15, or to detect a doubling of fly activity
(CV = 0.25) then just 6 fly ribbons, 5 spot cards, or 3 baited traps
are needed. With no published action thresholds for dairies or other
cattle facilities, Gerry et al (2011) used the highest mean fly or spot
count recorded for each monitoring method as a surrogate for the
unknown action threshold. If the true action threshold is lower than
the highest recorded mean count, then the number of traps needed
may be higher than the number reported in this study.
While the methods described above can also guide immature
sampling effort (number of samples needed) for routine monitoring
of fly activity at any one site, substantial aggregation of immature
flies within developmental substrate suggests that optimum sample
size is more appropriately derived with variance estimated using
Taylor’s power law (Stafford and Bay 1994). After sampling third-
instar house fly larvae from poultry feces in shallow-pit caged-layer
houses, Stafford and Bay (1994) determine that to detect a doubling
of the immature fly density would require 64–145 separate sub-
strate samples depending upon the mean density per sample at each
sampling site. To reduce the high degree of variance in immature fly
density among small substrate samples (Schoenburg and Little 1966,
Stafford and Bay 1994), several small substrate samples may be
combined into larger pooled samples prior to analysis (e.g., Mullens
et al. 1996). This may reduce the total number of samples that must
be processed to achieve the desired level of precision.
Immature sampling is more commonly used to compare imma-
ture fly density among sites or developmental substrates. When com-
paring different sites, the sampling effort needed to achieve a desired
level of precision in mean density will vary by site (Southwood 1978).
Furthermore, the optimum sample size needed for each site or sub-
strate is typically unknown when sampling is initiated. Because the
time to process each immature sample is typically much greater than
the time needed to obtain the sample, processing only the number of
samples from each site that are needed to achieve the predetermined
level of precision will save considerable time. This can be accom-
plished following an iterative method outlined in Schoenburg and
Little (1966) by obtaining a larger than needed number of samples
from each site, sequentially processing each sample, recalculating the
sample mean and variance after each sample (or number of samples)
is processed, and applying these recalculated values to determine the
level of precision achieved to that point. Precision is determined by
the confidence interval for and percent of deviation from the sample
mean. If the desired precision has not been met, additional samples
are processed. Upon reaching the desired level of precision in the
sample mean for that site, all remaining unprocessed samples can
be discarded. Thus, while the number of samples processed for each
site or substrate will be different, the sample mean for each site or
substrate is determined to the same level of precision.
Discussion
House fly behaviors such as flight are influenced by diel variation
and environmental condition (Dakshinamurty 1948, Parker 1962,
Pickens et al. 1967, Zahn and Gerry 2020) so that even when fly
density is unchanged, the frequency of behaviors (fly activity) will
vary as time of day or environmental conditions change (e.g., Lysyk
and Axtell 1985). The impact of time and environment is most acute
for instantaneous or near-instantaneous measures of house fly ac-
tivity such as the Scudder grid count, visual observations, station
counts, or the moving sticky tape method. When compared dir-
ectly, instantaneous measures of fly activity have been unrelated to
Journal of Economic Entomology, 2020, Vol. 113, No. 6
fly density (Pickens et al. 1972, Beck and Turner 1985) probably
as a result of the considerable influence of both time and the en-
vironment on fly behavior at any single point in time. Measures
of house fly activity with sampling times of <24 h were also un-
related to fly density (Pickens et al. 1972, Beck and Turner 1985)
likely for the same reason. Increasing the sampling period to ≥24 h
improved the relationship of fly activity estimates to fly density for
all monitoring methods. While monitoring methods are intended to
estimate fly activity, rather than fly density as discussed at the be-
ginning of this review, useful monitoring methods are still expected
to have a relationship to fly density. For example, fly density ac-
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counted for 54–60% of the fly activity estimate in poultry houses de-
pending upon the monitoring method used (Lysyk and Axtell 1985).
Monitoring methods used for longer sampling periods are expected
to provide a better estimate of overall fly activity because flies (or
spots) accumulate over a range of environmental conditions experi-
enced by the fly population being sampled (Lysyk and Moon 1994).
Instantaneous or short-term measures of fly activity may still be
useful to record change in fly activity when it can be reasonably as-
sumed that either fly density or frequency of fly behavior is relatively
unchanged to allow for interpretation of the estimated fly activity.
Sampling repeatedly throughout a day for example might provide
insight into environmental impacts on fly behavior (e.g., Zahn and
Gerry 2020). And studies that perform sampling at the same time
each day are attempting to reduce the environmental impact on fly
behavior so that changes in fly density can be examined. However,
the impacts of time and environment on fly behavior are complex
(Zahn and Gerry 2020) as suggested by the lack of correlation be-
tween fly density and instantaneous measures of fly activity where
this has been examined (Pickens et al. 1972, Beck and Turner 1985).
When all fly monitoring studies are considered together, regard-
less of facility type, fly activity estimates were related to fly density
when sampling periods were ≥24 h for fly ribbons, ≥2 d for baited
jug traps, or 7 d for spot cards (Pickens et al. 1972, Rutz and Axtell
1979, Lysyk and Axtell 1986, Stafford et al. 1988, Geden 1999,
Gerry et al. 2011). The magnitude and precision of fly activity esti-
mates is related to the design and placement of monitoring devices.
Fly activity estimates were higher for monitoring devices placed on
or near roof rafters (Rutz and Axtell 1979, 1981; Lysyk and Axtell
1985, 1986; Stafford et al. 1988), where house flies typically accu-
mulate at night (Anderson and Poorbaugh 1964). However, high
daily temperature can negatively impact fly capture near the roof
at least for spot cards (Lysyk and Axtell 1985). Also, spot cards or
sticky traps placed at downwind locations in ventilated housing will
outperform those placed at upwind locations (Geden et al. 1999).
House fly activity also varies along the long axis of a poultry house
(Willson and Mulla 1975, Burg and Axtell 1984) suggesting moni-
toring devices should be distributed along the long axis for any
animal housing structure.
Devices that require capture of flies on sticky surfaces (fly rib-
bons and sticky traps) are unsatisfactory in dusty conditions (Pickens
et al. 1967, Rutz and Axtell 1981, Kaufman et al. 2001, Gerry et al.
2011) and fly ribbons perform poorly outdoors particularly in direct
sunlight and high wind (Anderson and Poorbaugh 1965, Gerry et al.
2011). Traps baited with liquid attractants are generally unsuitable
for fly monitoring due to the lack of specificity for house flies and dif-
ficulty of identifying flies captured in the liquid bait. Baited jug traps
(or other devices using insecticidal bait) can rectify the problem of
separating flies from a liquid attractant, but these traps are unsuitable
for a long-term fly management program since any decrease in suscep-
tibility of the flies to the offered insecticide will confound fly activity
estimates over time (discussed in Geden 2005, Gerry et al. 2011).
Journal of Economic Entomology, 2020, Vol. 113, No. 6
Baited jug traps (and similar devices) are, however, well-suited for
short-term monitoring studies where fly susceptibility to the insecti-
cidal bait is unlikely to change. Baited jug traps can also be used
in dusty or outdoor environments where other monitoring devices
perform poorly.
Spot cards are perhaps the simplest monitoring method to use
since they are easily handled and fly identification is not required.
But spot cards provide only an overall index of pest fly activity, since
several fly species will deposit spots onto these cards and the spots
cannot be distinguished to fly species (Axtell 1970a, Pickens et al.
1972, Lysyk and Axtell 1986). However, if the purpose for moni-
toring flies is to reduce fly nuisance, then the lack of specificity to fly
species may be less important since fly species that commonly de-
posit fly spots are also those that can cause nuisance. Spot cards may
overestimate fly activity relative to other monitoring methods since
individual flies may deposit more than one spot onto a card while
methods that trap flies will count each fly only once. And overesti-
mation might increase at higher temperatures if spot deposition rate
is related to temperature. Spot cards will be damaged by water and
may be impacted by dust (Rutz and Axtell 1981) so are best used in-
doors and away from animal waterers or misting devices.
Fly ribbons allow for simultaneous monitoring of several pest
fly species (Anderson and Poorbaugh 1964; Axtell 1970a,b; Legner
et al. 1973; Williams and Westby 1979; Lysyk and Axtell 1986;
Gerry et al. 2011), though a species interaction may occur as the fly
ribbon begins to fill with flies (Axtell 1970a). Sticky cards or other
sticky traps are expected to have the same benefits and drawbacks
as fly ribbons, though sticky cards may be much easier to use than
most other sticky trap designs (Black and Krafsur 1985, Hogsette
et al. 1993, Geden et al. 1999). In outdoor locations, baited traps
and rigid sticky traps are probably best as they can more effectively
withstand the outdoor environment, though dusty conditions and
removal of captured flies by birds can negatively impact the use of
sticky traps (Kaufman et al. 2001, Gerry et al. 2011).
Fly management programs require action thresholds as a target
for fly control efforts. Currently, there are few action thresholds pub-
lished for guidance, and these thresholds are subjectively derived and
may be difficult to defend. A way forward may be to empirically
determine action thresholds for individual facilities by comparing fly
activity with presence or absence of nuisance complaints. This would
require ongoing monitoring of house fly activity using an appro-
priate monitoring method for evaluation against a future nuisance
complaint. An empirically determined action threshold will differ for
each facility according to site-specific factors to include the design
and operation of the facility, characteristics of the surrounding en-
vironment, distance to nearby homes and schools, and the tolerance
for flies of people in the surrounding area. In addition, the action
threshold will necessarily differ by fly monitoring method selected
and by the specific placement of individual traps or devices used.
However, as more data on fly activity related to nuisance becomes
available, it is possible that some consensus on action thresholds
may be found at least among methods for monitoring fly activity at
animal facilities with common design characteristics.
Additional needs include efforts to standardize trapping method-
ology, including the design, placement, and number of monitoring
devices to be used by type of animal facility. Research studies to
date have primarily focused on caged-layer poultry, and studies in
other animal facilities are needed. While fly activity estimates (and
action thresholds) may not be directly comparable across facilities
due to facility-specific differences in fly distribution and to differ-
ences in fly capture rates by design and placement of monitoring
2577
devices, commonalities in monitoring outcomes among facilities can
improve monitoring methods used across animal facilities.
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