BIOLOGICAL CONTROL 13, 16–24 (1998)
ARTICLE NO. BC980632
Biological Control: An Important Component in Integrated Management
of Musca domestica (Diptera: Muscidae) in Caged-Layer
Poultry Houses in Buenos Aires, Argentina
Diana C. Crespo,*,† Roberto E. Lecuona,* and Jerome A. Hogsette‡
*Instituo de Microbiologı́a y Zoologı́a Agrı́cola (IMYZA), CICA, INTA C.C. 25 (1712) Castelar, Buenos Aires, Argentina; †CONICET,
Buenos Aires, Argentina; and ‡USDA, Center for Medical, Agricultural, and Veterinary Entomology, Gainesville, Florida
E-mail: afn15308@freenet.ufl.org
Received May 15, 1997; accepted April 8, 1998
An Integrated Management program to control
Musca domestica L. was developed by evaluating cul-
tural 1 chemical control, cultural 1 biological control
1 granular bait (IMP 1), and cultural 1 chemical 1
biological control 1 granular bait (IMP 2) in poultry
houses in Argentina. Adult fly density was estimated by
a modified Scudder-grid method. Percentage of house
fly parasitism was estimated by direct and indirect
methods. Spalangia endius Walker and Muscidifurax
raptor Girault & Sanders were released at a rate of 10
parasitoids/hen/week. Complete absence of control
practices allowed a sustained growth of the fly popula-
tion. When larvicides and adulticides were used, the
number of adult flies decreased but parasitism was
adversely affected. When insecticide use was de-
creased, the number of flies increased. Cultural 1
biological control 1 granular bait (IMP 1) produced a
reduction of adults from 14 to 3 flies/grid. Where
cultural 1 chemical 1 biological control 1 granular
bait (IMP 2) was employed, the adult number was
reduced from 40 flies/grid to 1 fly/grid. IMP 2 was
efficient for reducing the number of larval develop-
ment sites in the manure. Percentage parasitism ex-
ceeded 90% in both IMP 1 and IMP 2 treatments. IMP 1
and IMP 2 treatments both allowed for maximum
control of house flies with minimal use of pesticides.
With IMP 2, the potential effectiveness of the biological
control agents was maximized by reducing adult fly
populations with pesticide sprays, then terminating
spray applications before parasitoids were released.
Thus, IMP 2 would be most suitable for farms which
have large adult house fly populations at the initiation
of the treatment. The only pesticide used in IMP 1 was
a granular bait. Thus, IMP 1 would be most efficient in
situations with low initial adult house fly populations
and small amounts of manure to ensure the success of
the cultural strategy. r 1998 Academic Press
Key Words: Musca domestica; biological control; aug-
mentative release; Spalangia endius; Muscidifurax rap-
tor; IMP; poultry houses.
1049-9644/98 $25.00
Copyright r 1998 by Academic Press
All rights of reproduction in any form reserved.
INTRODUCTION
Accumulated manure in commercial caged-layer
houses is a potential source of disease organisms
harmful both to confined hens and to humans (Anony-
mous, 1975). Because there are arthropods with a high
capacity for dissemination of metaxenic (Del Ponte,
1958) and other types of important diseases, this can
pose a potentially serious health problem. Among dip-
teran species, Musca domestica L. has an important
role as a disease vector (Aberg-Cobo et al., 1959). The
behavior of this pest is typically synanthropic and,
because of its high reproductive rate and ability to
prosper in a wide range of environments, it pullulates
throughout the entire year.
Oliva (personal communication) estimated that in
Argentina the annual cost of house fly control on a
50,000-bird poultry farm costs approximately $10,000
U.S. There are 16 million hens in the country (Schang,
1991), half of which are on poultry farms where in-
secticides are routinely used for house fly control.
This represents an annual cost of approximately
$1,600,000 U.S.
Besides being costly, insecticidal control of the house
fly has many serious drawbacks. If used improperly,
insecticides can produce intoxications in animals and
humans, contaminate feed and water, and destroy the
biological control agents of flies. Likewise, flies possess
a strong genetic capability for developing resistance to
the insecticides used by farmers (Wang and Plapp,
1980; Roush and Wright, 1986; Meyer and Georghiou,
1987; Shen and Plapp, 1990; Cluck et al., 1990; Kence
and Kence, 1992; Oi et al., 1992).
In recent years, Integrated Management Practices
(IMP) for house flies have become more widely used.
Combinations of different control alternatives, such as
biological, cultural, or chemical control, provide interest-
ing options for regulating fly populations without caus-
ing environmental damages of any importance (Axtell,
16
 INTEGRATED MANAGEMENT OF M. domestica IN POULTRY HOUSES 17

1986; Morgan et al., 1975b; Patterson and Ripa, 1982;
DeVaney et al., 1990).
Scientists in several countries, including the U.S.
and Canada, have been successful in using parasitoids
to control the house fly. Presently, this methodology is
applicable at the commercial level (Morgan et al.,
1975a,b; Axtell, 1986). In Argentina, research has been
done only to identify the natural enemies of M. domes-
tica, notably pteromalid (Hymenoptera) parasitoids,
which are effective against the pupae of the house fly
and other related Diptera (De Santis and De Sureda,
1988).
The purpose of our work was to develop an Integrated
Management Program for house fly control using pupal
parasitoids as a biological control component, optimiz-
ing hygienic and economical aspects and minimizing
environmental contamination risks. Fly control would
be considered acceptable when the population densities
in poultry houses were #10 flies/grid when measured
with Scudder grids. To this purpose, different control
strategies were evaluated in commercial laying houses
located near Buenos Aires, Argentina.
MATERIALS AND METHODS
The experiments were performed on four caged-layer
farms located in Pilar and General Rodriguez, Buenos
Aires Province, Argentina, during 1992 and 1993. Ex-
cept in treatment a, the untreated control, selected
pesticides were used to reduce adult fly populations. In
treatments that included a biological control compo-
nent, adulticides were used to give the pupal parasi-
toids numerical superiority over the house flies and to
help reduce the overall fly population. Treatments are
described in detail below and summarized in Table 1.
(a) Untreated control. Two houses with about 6,000
hens each served as untreated controls in which no
house fly management techniques were performed.
When the evaluations began, manure had accumulated
under the cages for 2 months. This was similar for
treatments b and d as well.
(b) Cultural plus chemical control. This treatment
was applied to a house with 10,000 hens where water
leaks were eliminated. Initially, manure was treated
with quicklime (calcium oxide) and slaked lime (cal-
cium hydroxide) mixed in a proportion of 3:1, respec-
tively. The amount used per m2 was dependent upon
the size of the larval foci and moisture content of the
manure. Localized lime treatments were applied on
three occasions to wet spots in the manure, and general
repairs were made in various parts of the house. DDVP
(Nuvan 100 EC (100 g/liter AI), Ciba-Geigy Buenos
Aires, 250 cc/20 liter of water) was applied once and
azamethiphos (Alfacron 10% (AI), Ciba-Geigy Buenos
Aires, 250 cc/250 liter of water) was applied as a
paint-on every 2 weeks for control of adult flies. Larvae
were controlled by feeding cyromazine (Larvadex 1%
(AI) premix, Ciba-Geigy Buenos Aires, 500 g/per ton of
poultry feed) for five consecutive weeks, accompanied
by three topical applications of cyromazine (Neporex
50% (AI), Ciba-Geigy Buenos Aires, 1 g/m2 ) to manure
ad libitum. Snip, which contains a sexual attractant
and a toxicant (z-9-tricosene with azamethiphos 1%
(AI), Ciba Geigy, Buenos Aires), was applied as a
granular bait at the rate of 2 kg/house/month as an
adulticide. At this farm, all mentioned chemicals were
used for the first time during these evaluations.
(c) Cultural plus biological control plus granular bait
(IMP 1). This treatment was applied in six houses,
each with 10,000 caged-layers. Houses were free of
manure when birds were housed, and lime had been
applied to the floor in manure collection areas under
the cages to help dry the manure. Lime was also
applied ad libitum on two occasions to wet spots in the
manure. Spalangia endius (Walker) and Muscidifurax
raptor (Girault & Sanders) were released weekly at the
rate of 10 parasitoids/hen/week (50:50 ratio) starting at
the beginning of the evaluation (January). Releases in
each house were made by placing approximately 2,500
parasitized fly pupae (30 g) in each of 40 paper bags
(18 3 10 cm), which were then suspended from the
floors of the cages as close to the aisle as possible. Bags

TABLE 1
Description of Poultry Farms and Treatments and Components Assigned to Each

Farm Number
number of houses Treatment number and type Components

1 2 houses (a) Control None

2 1 house (b) Cultural plus chemical Moisture control 1 lime 1 nuvan 1 alfacron 1 larvadex
1 neporex 1 Snip
3 6 houses (c) Cultural plus biological control plus granular bait Moisture control 1 lime 1 Snip 1 parasitic wasps
4 2 houses (d) Cultural plus chemical plus biological control plus Moisture control 1 lime 1 nuvan 1 Snip 1 parasitic
granular bait wasps

Note. Larvadex, cyromazine 1% premix; neporex, cyromazine 50% topical; nuvan, DDVP 100% AI; alfacron, azamethiphos 10% paint; Snip,
azamethiphos 1% AI (with z-9-tricosene) granular bait; parasitic wasps, Spalangia endius (Walker) and Muscidifurax raptor (Girault and
Sanders).
18 CRESPO, LECUONA, AND HOGSETTE
were spaced 14 m apart and the tops opened slightly to
allow parasitoids to exit. Parasitoid emergence began
soon after placement of bags in houses and continued
for approximately 1.5 days. Snip granular bait was
applied as in treatment b above.
(d) Cultural plus chemical plus biological control
plus granular bait (IMP 2). This treatment was ap-
plied in two houses containing 10,000 hens each. The
cultural management techniques (lime) were similar to
those for treatment b. DDVP was sprayed once for
control of adults only in localized places and discontin-
ued after parasitoids were released. Parasitoids were
released in the same way as in treatment c, but no
releases were made until the 7th week of the evalua-
tion. Snip granular bait was applied as in treatment b
above.
Density estimates of house fly adults. The modified
Scudder grid method (Murvosh and Thaggard, 1966)
was used to estimate adult fly populations in the
treated and control houses. Estimates were made
weekly from December 12, 1992 to May 15, 1993. Fly
counts were made in areas of high fly density through-
out the houses by placing a grid (45 3 45 cm) on the
manure surface and counting the flies landing on it
during a 1-min period. The average number of flies per
grid for each treatment was determined by selecting
five grids with the greatest number of flies within a
variance range of ,5%. Fly control was considered to be
at an optimal level when the population density in the
houses was #10 flies/grid.
Rearing of parasitic wasps. Techniques for rearing
indigenous S. endius and M. raptor released during the
FIG. 1. Adult house fly density in control treatment a.
evaluations were similar to those described by Morgan
et al. (1978). Rearing rooms were maintained at 27 6
1°C and 70 6 10% RH, with a 24-h scotophase. Parasi-
toids were reared in the pupae of house flies produced
in an artificial diet (adapted from Morgan, 1986).
Percentage parasitism. Percentage parasitism of the
house fly was estimated by an indirect method using
sentinel pupae (Rutz and Axtell, 1981) and a direct
method where pupae were extracted from substrates in
the poultry houses. For the indirect method, colony-
reared house fly pupae ,24 h old were placed in
open-mesh bags (10 3 10 cm) at the rate of 30 pupae
per bag. Bags with pupae were placed in the manure
under the cages at the rate of 30 bags per house per
week. Pupae were held in the laboratory for 30 days for
the emergence of parasitoids, at which time the percent-
age parasitism was calculated. For the direct method,
500 house fly pupae per house per week were collected
at random and held for 30 days at ambient temperature
in the laboratory for the emergence of parasitoids.
Percentage parasitism was calculated with the intact
puparia method of Petersen (1986) using the formula:
percentage parasitism 5 ((emerged 1 aborted
parasitoids) 4 intact puparia) 3 100.
RESULTS AND DISCUSSION
(a) Untreated control. Mean M. domestica popula-
tion in the controls (a) increased from 44 adults/grid at
the beginning of the experiments to 192 adults/grid 15
weeks later (March 18, 1993) (Fig. 1). The trend of
increasing house fly populations with the progression
 INTEGRATED MANAGEMENT OF M. domestica IN POULTRY HOUSES
of the season can be seen, particularly from December
until March. We attributed this population increase to
the complete absence of fly control practices.
(b) Cultural plus chemical control. At first, when
larvicide (cyromazine) and adulticides (such as baits
with sexual lures and DDVP sprays) were used to-
gether in the cultural plus chemical control treatment
(b) (Fig. 2), fly populations decreased from 41 to 6
flies/grid after 5 weeks. Beginning January 23, 1993,
cyromazine was discontinued for 3 weeks and the
number of house fly adults increased from 6 to 12
flies/grid. A second application of cyromazine was made,
which reduced the number of flies to 2/grid. When
cyromazine was again discontinued (March 13, 1993),
fly density increased slightly, but stayed ,9 flies/grid.
This phenomenon, reported also by Morgan et al.
(1975a), was observed on both occasions when cyroma-
zine was used and then discontinued.
It is worthy to note that genetic resistance to cyroma-
zine and the adulticides was not observed in these
house fly populations, possibly because this was the
first time these products were used on this poultry
farm. The combined use of both strategies (cultural
plus chemical) succeeded in maintaining the adult fly
population at tolerable levels (Fig. 2).
(c) Cultural plus biological control plus granular bait
(IMP 1). The cultural plus biological control treat-
ment plus granular bait (c) began with an initial adult
population of 14 flies/grid. However, adult fly popula-
tions decreased from 14 to 4 flies/grid after the first 2
weeks of parasitoid releases and were maintained at
FIG. 2. Adult house fly density in cultural plus chemical treatment b.
19
about 3 flies/grid for the next 8 weeks (Fig. 3). The
overall lime application provided a dry base for the
initial layers of manure, and the two localized applica-
tions eliminated fly breeding in wet spots.
Treatments were discontinued at the beginning of
April because hens had become infected with fowl
typhoid (Salmonella gallinarum), which causes ex-
tremely wet droppings. This stresses the importance of
monitoring the health and sanitary condition of con-
fined animals, especially for diseases which cause
diarrhea. With the combined strategy of cultural plus
biological control plus granular bait, the pest popula-
tion was maintained within acceptable levels (,10
flies/grid) until treatments were discontinued. How-
ever, the effectiveness of the last parasitoids released
was probably reduced because of the increase in ma-
nure moisture caused by the diarrhea of the hens.
(d) Cultural plus biological plus chemical control
(IMP 2). In houses receiving the IMP treatment (d),
the mean number of adult flies was reduced from 40 to
17/grid during the first 7 weeks (Fig. 4). Lime and
adulticide reduced the number of larval foci, substan-
tially improving the general condition of the houses.
Weekly releases of S. endius and M. raptor began on
January 30, 1993 (Fig. 4), and in 5 weeks adult house
fly population density was reduced from 17 to 1/grid.
Good control was maintained throughout the study,
with adult populations never exceeding 4 flies/grid.
Percentage parasitism. Percentage parasitism of
house fly pupae scored either by the direct or the
indirect method was very similar in the controls (a) and
20 CRESPO, LECUONA, AND HOGSETTE

FIG. 3. Adult house fly density in cultural plus biological treatment plus granular bait (IMP 1) (c).

the cultural plus chemical treatment (b); but, weekly
average values were lower when estimated by the
direct method compared with the indirect method
(Figs. 5 and 6). In contrast, the percentage parasitism
and the weekly average values calculated by either of
the two sampling methods were similar in both the IMP
1 (c) and the IMP 2 (d) treatments (Figs. 7 and 8).
When percentage parasitism is low, as it was in
treatments a and b, indirect sampling methods tend to
yield higher parasitism values than direct methods.
Direct methods are based on making collections of
mixed-age, naturally occurring, and possibly parasit-
ized, house fly pupae that may be clustered or scattered
throughout the study site. Finding enough pupae to

FIG. 4. Adult house fly density in cultural plus biological plus chemical control (IMP 2) (d).
 INTEGRATED MANAGEMENT OF M. domestica IN POULTRY HOUSES 21

FIG. 5. Comparison of percentage parasitism of house fly pupae in the control treatment (a) using the indirect and direct sampling
methods.

sample can at times be difficult (Rutz, 1986). Indirect
methods are based on exposing concentrations of
known-age, laboratory-reared house fly pupae in selected
sampling sites to attack by parasitoids; but this may
result in a high rate of parasitism produced by a small
number of wasps. In treatments c and d, where there
was little or no pesticide used for larval control, back-
ground larval populations were developing to the pupal
stage and being heavily parasitized by the large parasi-
toid populations being maintained during the study.

FIG. 6. Comparison of percentage parasitism of house fly pupae in the cultural plus chemical treatment (b) using the indirect and direct
sampling methods.
22 CRESPO, LECUONA, AND HOGSETTE

FIG. 7. Comparison of percentage parasitism of house fly pupae in the cultural plus biological treatment (c) using the indirect and direct
sampling methods.

Thus, results from direct and indirect methods were
very similar. However, when a percentage parasitism
.90% was observed 5 to 6 weeks after the initial
release, superparasitism was observed (dudding effect)
and direct sampling was difficult to perform because of
the scarce number of host pupae. Both the direct and
indirect method of pupal sampling have advantages
and disadvantages (Rutz, 1986), but Petersen and
Watson (1992) recommend that both methods be used
together when possible.
Percentage parasitism was almost 100% for several
weeks in treatments c and d (Figs. 7 and 8), and both

FIG. 8. Comparison of percentage parasitism of house fly pupae in the IMP treatment (d) using the indirect and direct sampling methods.
 INTEGRATED MANAGEMENT OF M. domestica IN POULTRY HOUSES
parasitoid species were recovered in almost equal num-
bers. Morgan et al. (1981) recorded similar parasitism
levels when releasing S. endius and M. raptor together;
however, they recovered very few M. raptor. Their
parasitoids were released from 15 stations outside the
poultry houses or from eight stations inside the poultry
houses. M. raptor apparently did not disperse from
these stations to fly breeding sites in the manure
(Morgan et al., 1981). In our study, parasitoids were
released at 14-meter intervals inside the houses, which
provided a very uniform distribution of parasitoids as
they emerged.
Based on local conditions, we determined that a
successful fly control program should maintain average
fly densities at #10 flies/Scudder grid inside the poul-
try houses. Other fly control programs have been based
on fly density estimates derived from Scudder grids
(Morgan et al., 1981), spot cards (Rutz and Axtell,
1979), sticky tapes (Turner et al., 1992), and baited jug
traps (Burg and Axtell, 1984). These devices provide
visible estimates which can be used by poultry produc-
ers to make subsequent control decisions.
Eventually, treatments b, c, and d all reduced density
estimates to #10 flies/grid (Figs. 2–4). However, our
treatment of choice was the cultural plus chemical plus
biological control (d). With this treatment, attempts
were made to eliminate wet spots in the manure with
lime and reduce adult fly populations with nuvan
before parasitoids were released. Control of wet spots
continued during the parasitoid release program, and
Snip provided additional adult fly control without
adversely affecting the parasitoids. The cultural plus
biological control treatment (c) looked very promising,
despite the hens’ disease problems, and employed Snip
as the only pesticide component. Percentage parasitism
peaked earlier with this treatment (Fig. 7) than it did
with treatment d (Fig. 8); however, initial fly popula-
tions were considerably lower in the houses used for
treatment c (Fig. 3) compared with those used for
treatment d (Fig. 4). The cultural plus chemical treat-
ment (b) stabilized fly populations below the optimal
level after 9 to 10 weeks (Fig. 2), but alfacron and snip
were routinely applied for adult control until the end of
the study.
In the cultural plus chemical treatment (b), each
time chemical products were applied inside the poultry
houses to reduce the adult fly population, the percent-
age parasitism was adversely affected. The DDVP
application abruptly reduced the percentage parasit-
ism from 34% on the application date to 10% 2 weeks
later (Fig. 6). This was not surprising, because DDVP
has proved to be moderately to extremely toxic to some
strains of Spalangia and Muscidifurax (Rutz and Scott,
1990). Also, immediately after the larvicide application
the percentage of parasitism underwent a second de-
crease from 38 to 18% in 4 weeks. This could be
23
ascribed to the cyromazine, but under our working
conditions these reductions were impossible to explain.
However, in laboratory experiments Klunker (1991)
demonstrated that fly pupae exposed to cyromazine
and larviform fly pupae resulting from exposure of fly
larvae to cyromazine have a significant adverse effect
on the immature stages of S. nigroaenea (Curtis), S.
cameroni (Perkins), and M. raptor.
An important first component of treatments b, c, and
d was moisture control, which included drying wet
manure areas with lime, and repair and maintenance
of watering systems and poultry housing. House flies
develop well in poultry manure with a moisture range
of 60 to 75% (Miller et al., 1974), but they have been
recovered in manure with moisture levels from 37 to
80% (Stafford and Bay, 1987). Dry manure is not only
important for the control of flies and odors, but it is
critical for maximum performance of parasitoids and
other natural enemies (Morgan et al., 1981).
CONCLUSIONS
The cultural plus biological plus chemical control
(IMP 2) (d) and cultural plus biological control plus
granular bait (IMP 1) (c) treatments were the most
appropriate strategies for successfully reducing house
fly populations to low levels and maintaining them at
low levels over time.
The IMP 2 treatment (d) would be better for those
cases in which initial house fly populations per house
are high, because the cultural and chemical strategies
used in the IMP treatment permit the number of adult
flies to be reduced before the biological control compo-
nent is used.
The IMP 1 treatment (c) would be better for those
situations with a low initial number of adult flies
because no pesticide sprays are used to reduce adult fly
populations. IMP 1 would also be better in situations
with a low initial quantity of manure to ensure the
success of the cultural strategy.
In all cases, it is important to be aware of the hens’
level of health and the structural condition of the
poultry houses. Neglect of these factors can result in
increased manure moisture, the development of larval
foci and the subsequent increase of the population
density of adult flies. In both treatments where para-
sitic wasps were used, the biological control potential
was maximized by using selected pesticides to reduce
adult fly populations without contaminating areas occu-
pied by pupal parasitoids.
ACKNOWLEDGMENTS
We thank Dr. Renato Ripa S. (INIA, La Cruz, Chile) for many
valuable suggestions and for his critical reading of the manuscript.
We also are grateful to Dr. Pascual Franzone and Mrs. Estela Favret
for their help with the translation.
24 CRESPO, LECUONA, AND HOGSETTE
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