Spectral Sensitivity of Stable, Face, and Horn Flies and Behavioral

Responses of Stable Flies to Visual Traps (Diptera: Muscidae)1

H. R. AGEE' AND R. S. PATIERSON'
Agricultural Research Service. U.S. Department of Agriculture, Gainesville, Florida 32604

Environ. Entomo\. 12: 1823-1828 (1983)

ABSTRACT Compound eyes of laboratory-reared and field-collected stable and horn flies, and lab-
oratory-reared face flies have peak visual sensitivities from 360 to 490 nm. Hom and stable tlies have
a visual sensitivity plateau at 625 nm that was not detected in laboratory-reared face flies. Clear fiber
glass and ultraviolet reflectors were the most efficient materials for trapping stable flies. Traps were
more efficient when exposed to full sunlight which enhances reflection of ultraviolet light. In compar-
ative tests, materials with poor ultraviolet refl.ection were less effective for capturing stable flies.

Fly populations have been monitored for many years
with various types of traps. Hansen (1951), one of the
first to use a visual-response trap in muscoid flies, used
a black-painted board to attract stable flies, Stomoxys
calcitrans L. It was assumed that the stable fly easily
detected the board because of its visual contrast to the
surrounding white-sand beaches. Conversely, La-
Brecque et al. (1972) used boards painted white to mon-
itor stable fly populations in dark, unpainted feed bunkers.
Again, it was assumed that the black-white contrast at-
tracted the flies. Pospisil and Zdarek (1965) felt that
stable flies were attracted to contrasting objects rathcr
than to color, a conclusion that is supported by the ob-
servations of both Hansen and LaBrecque et al. Yet it
has been reported that dark-colored cattle and dark areas
on cattle consistently have higher numbers of stable and
hom flies, Haematobia irritans L., than lighter animals
or lighter areas on animals in the same herd (Parr 1962,
Bidgood 1980). Gatehouse and Lewis (1973) and Pos-
pisil and Zdarek (1965) felt that flies landed preferen-
tially on low-reflectance surfaces rather than selection
of surfaces of specific colors. Patterson and Juma (1980)
observed that certain animals, regardless of color or size,
consistently had higher stable fly counts, even though
all animals appeared similar in color and size to the
casual observer. Their observation suggests that other
factors also may playa role in host preference.
Williams (1973) found that, when he applied the ad-
hesive Tack Trap to the clear fiber glass Alsynite instead
of the standard white-painted plywood used by La-
Brecque et al. (1972), more stable flies were captured.
Pickens et al. (1977) found that white-painted pyramid
traps were very attractive to stable and face flies, Musca
aUTlImna[is De Geer. Therefore, traps designed using
Alsynite for stable flies and white paint for face flies
were effcctive attractants. In fact, these traps were so
efficient that area populations were reduced (Meifert et
al. 1978).
When we tested brands of fiber glass other than AI-
synite, some were unattractive to stable flies. Therefore,
IMention of a commercial or proprietary product does not constitute
an endorsement by the USDA. Received ror publication 28 April 1983:
llccepted 16 August 1983.
~Insecl AUraelanls. Behavior. and Basic Biology Research Labo·
rJto~.
Insecls Affecting Man and Animals Research Laboratory.
the primary objective of the studies reported hcre was
to determine whether a correlation existed betwcen the
spectral sensitivities of the compound eyes of face, horn,
and stable flies and the spectral rcflcctance of various
surfaces. Since face flies arc not found in Florida, the
attractiveness of various surfaces was conducted with
hom and stable flies. Both specics are biting, diurnal
livestock pests attracted to light--cspecially ultraviolet
(UV) light when placed in a dark environment (Driggers
1971, Milleret al. 1979, Patterson and Miller 1982).
Thc secondary objective was to determine if any dif-
ference existed between the spectral sensitivities of lab-
oratory-reared and wild flies and what differences, if
any, might be caused by age or sex. Furthermore, since
it was known that Alsynite fiber glass was very attractive
to stable flies, we wanted to determine if the spectral
reflectancc of the fiber glass changed with age, usc, or
the addition of adhesive (Tack Trap) or insecticide (per-
methrin).
Materials and Methods
Electrophysiological Studies
The instrumentation and techniques used to measure
the visual sensitivity of the three species of flies were
similar to those used by Agee (1973). Monochromatic
light was produced by an equipment system consisting
of a quartz-halogen lamp, quartz-iconel neutral-density
filters, a light chopper, color filters for second-order
wavelengths, and a monochromator with settings of 5-
nm half-height band widths. A stimulator controlled the
light stimulus duration and rate via a small, solenoid-
operated metal vane. The flies were mounted on a test
platform inside a light-proof Faraday cage and posi-
tioned with the right eye in the stimulus light beam and
the left eye in the dark. Stainless-steel electrodes were
etched to fine points, and the recording electrode was
inserted ca. 20 J-lm into the dorsolateral region of the
right eye. The indifferent electrode was positioned sim-
ilarly in the left eye. A biological amplifier amplified
the voltage changes that occurred when the eye was
stimulated with IOO-msecpulses of monochromatic light
at wavelengths from 350 to 700 nm at a rate of one
pulse per 2.5 sec. A "criterion" response of 200 J-lV
was required as a standard. A 30-min dark adaptation
period was lIsed for all flies once we determined that

1823
1824 ENVIRONME"TAL ENTOMOLOGY Vol. 12, no. 6

thin layer of nondrying adhesive (Tack Trap). These

0.005 sticky panels were placed in a straight line ca. I m apart
along a pasture fence. Stable fly- or horn f1y-infcsted
cattle and horses were pastured within the fenced pas-
0.01
ture. After 24-h exposure periods. the panels were checked
"e••
....• for fly capture. and panels werc randomly rotated among
! positions. The species and sex of captured flies were
determined.

i
en
0.05
)"\I~.\.
Results and Discussion
E/ectrophysiological Studies

~
oJ
0.10
I ~ The spectral sensitivity curves for wild and labora-
tory-reared stable flies (male and female combined) are
'"
0
~
m \ shown in Fig. I and 2, respectively. Both sexes of lab-
oratory-reared (I to 6 days old) flies have major peaks
m_\
"~
II:

W
0.50
1.00
350 400 450 500 550 800 850
WAVELENGTH IN NANOMETERS
FIG 1. Mean spectral sensitivity of the wild stable fly, S.
ca/citrans L. (five of each sex, age unknown).
visual sensitivity did not increase after additional dark-
adaptation periods. The thermopile was calibrated by
Eppley Laboratory, Inc. (Newport, R.I.), in microvolts
per microwatts per cm' at three intensities relative to a
carbon filament standard lamp from the National Bureau
of Standards.
Spectral sensitivity tests were performed on labora-
tory-reared and field-collected stable and horn flies of
both sexes at selected ages. Only laboratory-reared face
flies were tested because they are not found in Florida,
and therefore were held in quarantine during the test
of visual sensitivity in the 450- to 500-nm region (peak
at 490 nm) as do the wild flies. A secondary peak occurs
in the UV at 360 and a plateau in the sensitivity curve
at 625 nm for all groups except the laboratory-reared
stable flies when I to 2 days old. Laboratory males and
females do not differ significantly in visual sensitivity,
and field-collected flies of both sexes are not signifi-
cantly different from thc laboratory-reared stable flics
(Mann-Whitney U-test[Sokal and Rohlf 1969]). Ballard
(1958) found that both sexes of stable fl ies responded
well to 365-, to 465-. and to 640-nm stimuli.
The spectral sensitivity curve of the face fly was sim-
ilar to that of the stable and horn flies, in that the com-
pound cyes of the IO-day-old face flics were most scnsitive
to 360-nm (UV) stimuli. with a broad peak centered on
0.005
0.01
"'Eu
....•

period. Each study was replicated 10 times, and spectral !

sensitivity tests were performed 0 to 3 h after a citrated en
bovine blood meal unless otherwise indicated. 3
::l

Spectral Reflectance of Trap Surfaces

!
en
0.05

!i:
Trap spectral reflectance was measured to determine
if a correlation existed between the spectral sensitivity ":i
IL
0.10

of each species and the light reflecting from the trap 0

surfaces. Reflectances of samples 2.5 by 2.0 cm were >-

measured with a Varian 634S recording reflectance "
II:
W
m_Cl ~~
spectrophotometer. The surfaces were scanned at wave- iii
.-.!!.~
lengths from 330 to 850 nm, with the monochromator 0.50 II-II !-~
slit width set at 1 mm and operated at a scan rate of 50
nmlmin to obtain maximum accuracy. Eastman white
(no. 6091) was the 100% reflectance standard. 1.00
350 400 450 500 550 800 850

Attractiveness of Trap Surfaces WAVELENGTH IN NANOMETERS

Various surfaces were field evaluated for attractive-
ness to both stable and horn flies. The material to be
evaluated was cut into a 30-cm' panel or placed on a
frame of that size. The test surfaces were coated with a
FIG. 2. Comparison of mean spectral sensitivity of labora-
tory-reared 3-day-old face flies, M. o/ilul1Illa/is De Geer (20
each), hom flies, H. irritans L. (10 each), and stable flies. S.
co/citrons L. (10 each). (Approximatelyequal numbersof each
sex in each group.)
December 1983 AGEE AND PATIERSON; FLY VISION AND TRAPPING 1825

490 nm and a plateau of sensitivity to 625-nm stimuli

0.005
on the declining red end of the curve (Fig. 2). Both
males and females responded similarly, which is in gen-
eral agreement with the recent face fly study of Pickens 0.01
(1983). ~
The compound eyes of male and female, 10-day-old, "
.....
laboratory-reared face flies were most sensitive to: (I) !
the 360-nm UV; (2) a broad peak centered on 490 nm;
and (3) a small plateau of sensitivity at 625 nm on the
declining side of the curve (Fig. 2 and 3). Goldsmith
and Fernandez (1966) also found that the dark-adapted
house fly, Musca domestica L., had a small peak of
i 0.05

0.10
visual sensitivity at 620 nm, and they concluded that it ~
was' 'caused by a transparency of the accessory screen-
~
ing pigment to long wavelengths."
>
Horn flies (laboratory-reared and field-collected) had
"1i!i5 .-.
a broad major peak of sensitivity at 490 nm, with a
second peak at 360 nm. Unlike stable flies, wild horn
flies did not have a small sensitivity plateau at 625 nm
(Fig. 4).
0.50
0-0
-
F_'

In laboratory-reared and field-collected horn flies there 1.00

were changes in sensitivity across the spectrum, de- 350 400 450 500 550 800 850
pending on the time lapsed between the last blood meal WAVELENGTH IN NANOMETERS
and the test period. Field-collected and laboratory-reared FIG. 4. Mean spectral sensitivity of wild hom flies, H.
horn flies that had not been fed a blood meal within 3 irritans L., (five males and females) measured 3 to 6 h after
to 6 h had reduced visual sensitivity and higher mortality blood meal.
than those provided a meal. All laboratory-reared flies
died within 20 h if not fed a blood meal.
response. Stable flies were more sensitive to 490-nm
All three species tested were most sensitive to 490-
stimuli. Pickens (1983) found three distinct photoreccp-
and 360-nm stimuli. Based on spectral sensitivity, it
tors for the face fly with sensitivity peaks at 350, 475
appears that face flies are more easily attracted to ma-
to 525, and 550 nm, respectively.
terials that reflect well at wavelengths of 360 nm, since
it was the wavelength that exhibited the greatest receptor
Spectral Reflectance of Trap Surfaces

0.005
. . Clear Alsynite is highly attractive to both sexes of
stable flies of mixed ages and shows an increase in re-
flectance in the 380 to 420-nm wavelengths. As Alsynite
weathers and turns slightly yellow (to the human eye)
0.01 -
(Fig. 5), its attractiveness is reduced 25% and the re-
"g
.....
~
flection on the UV end of the spectrum decreases with
! a shift to 450 nm. The reflectance of Alsynite covered
with either permethrin or Tack Trap had a similar spec-
tral reflectance pattern to the clear fiber glass (Fig. 5).
~ If a UV reflector such as HM-35 (Day-Glo Color
0.05
~ Corp., Cleveland, Ohio) is added to the surface of the
Alsynite, there is an increased reflection of wavelengths
~
~ 0.10 - -~ from 350 to 410 nm. Alsynite traps were tested after a
24-h exposure period in the field after they were coated
~ with dust and dead insects. The reflectance pattern was
>
c:J
F_ ~
• changed extensively (Fig. 58), with a reduction to 2%

! 0.50
.
.-.
0-0

•••••
reflectance at 390 to 420 nm and an increased reflec-
tance at wavelengths from 400 to 625 nm .
White Alsynite showed the greatest reflection at
wavelengths from 390 to 450 nm. Green Alsynite had

1.00
350 400 450 500
.
550 800 850
a peak reflection at 390 to 480 nm, but no sharp increase
occurred at these wavelengths as it did for the clear and
white (Fig. 5A). Of the colors listed in Table I, those
WAVELENGTH IN NANOMETERS itemized below clear Alsynite and light pink are rela-
FIG. 3. Mean spectral sensitivity of IO-day-old, laboratory- tively poor visual attractants. The light pink was unusual
reared face flies, M. aU/umnalis L. (10 of each sex). in that capture rate on it approached that of the gloss
1826 ENVIRONMENTAL ENTOMOLOGY Vol. 12, no. 6
......
......,...-
- ......,
..........•..
-..--,-.,..-"T.--
']. ~f
100

-- ------
1].-1 CLEAR --r
1/
eo

10
cu._ WE'TIERED -t
Ul

~1]J CLEAR C PERMETHR~ ~ r

1 ~
Ul

:~a: 10 ,.-/ CLEAR c TACK TRAP (NEW) ~tI

30
0
~
A 20
20

lQ

B 10

WAVELENGTH IN NANOMETERS o
350 400 450 500 550 600 eso 100 750 800 850

WAVELENGTH IN NANOMETERS

Flo. 5. (A) Spectral reflectance of trap surfaces used to capture hom flies, H. irritans L., and stable flies, S. falcitrans L.
(B) Spectral reflectance of Alsynite panels used to capture hom flies, H. irriwns L., and stable flies, S. ('alci/rans L.

white and clear Alsynite, The spectral reflectance char- Table 1. Comparison of the attractancy of selected materials
and colors for capturing stable nies (clear Alsynite captured 100%)
acteristics of white and light pink are quite similar for
wavelengths from 330 to 400 nm. Materials 'A CaplUre Color 'A: Capture
We found that most brands of nondrying adhesives
used in insect control programs masked reflectance of Carbon dioxide (CO2) 141 Clear Alsynite 100
the test material, making the surface unattractive to sta- White (gloss) 90
ble flies. Of the brands tested, only the commercial Alsynite + HM-35 140 Light pink 85
product Tack Trap did not alter the attractiveness of the Resins" 138 Lighl blue 63
White Alsyniteb 123 Light green 62
trap material (Fig. 58). Clear Alsynite 100 White (tlat) 61
Green Alsynite 57 Black 41
Attractiveness of Various Surfaces Alsynite (field)" 39 Brown 37
Plexiglas 35 Red 30
Alsynite + CO2 34 Yellow II
Of the 30 different surfaces field tested for attractancy Aluminum 29 Gold 10
to stable flies, the clear fiber glass, Alsynite (600 se- Plywood 14 Silver 0.2
ries), gave the most constant and successful results. Fur- Glass 3
thermore, this material is inexpensive, durable, and Saran wrap 2
Glass cloth 2
reusable with minimum cleaning effort. Therefore, its Polystyrene I
spectrum and trapping efficiency became the standard
by which the other products and colors were measured.
"Resin and fiber glass material used to make Alsynite.
Unpainted wood served as a second standard, because
bClear Alsynite performed bener than white Alsynite on cloudy days
flies often used wood buildings as a resting site but and in the shade.
would show no preference for it when placed alongside "A clear Alsynile panel after a 24-h exposure in field with aboUI four
a building where stable flies were resting. As shown in flies plus OIher insects and accumulated dust per square inch.
Table I, several other surfaces exhibited greater attrac-
tiveness during the tests than the clear Alsynite. The
most noticeable was white Alsynite that consistently ness may be a result of thermoregulation. For example,
outperformed the standard by ca. 23% during cool weather during cooler mornings and evenings, flies will congre-
(20 to 25°C) in bright sunlight. Some of the attractive- gate on white surfaces. presumably to thcrmoregulate
December 1983 AGEE AND PATIERSON;
body temperature. On warm or hot days. this behavior
was not noticeable. Our findings were consistent with
previous studies that clear Alsynite captured more flies
than white on cloudy. overcast, or hot days (Williams
1973. Ruff 1979. Bushman and Patterson 1981). White
and clear Alsynite have very similar reflectance curves
in the 360- to 4oo-nm range-ca. 15%; the reflectance
of white Alsynite gradually increases to 40% at 460 nm.
and clear Alsynite levels off at ca. 420 nm (Fig. 5A,B).
Green Alsynite attracted only 57% as many flies as the
clear (Table I), which was consistent with the results of
Williams (1973) and Ruff (1979), and exhibited only
21k reflectancy in the 360- to 420-nm range. Aluminum
attracted very few flies and was only 29% as attractive
to stable flies as the clear Alsynite standard (Table. I).
UV light is reflected by the 2-(2-hydroxy-5-methyl-
phenyl-benzolrizol) in Alsynite and is used at the rate
of 0.3 to 0.5%, by weight, in the resin (Reichhotd Inc.,
personal communication, technical representative). When
the resin was tested alone. it attracted more than one-
third as many flies as the clear standard. When the fiber
glass cloth was field evaluated. it appeared to repel sta-
ble flies. Almost 90% fewer flies were captured on the
fiber glass cloth than on the plywood standard. Since
many commercial products are made of similar UV-
absorber resins, we tested several to determine their ef-
fectiveness. All were more attractive to stable flies than
Alsynite (capturing 33 to 50% more) but were brittle
and shattered readily when subjected to stress. There-
fore. none were considered acceptable for field traps.
When the UV brightener HM-35 was coated on the
surface, stable fly catch increased 40%. The HM-35
brightner increased reflectance to 25% in the range from
350 to 400. or two times that of the clear Alsynite (Fib.
58). Since it was difficult to keep HM-35 on the fiber
glass under field conditions. it was impractical to treat
the panels to increase catch.
Dust and dead-fly buildup on trap surface reduced trap
efficiency. When these traps were evaluated in the field,
we found that. after 24 h, the traps collected only 39%
as many flies as in the first 24 h.
McCann and Arnett (1972) showed that the house fly,
M. domestica, had peaks of sensitivity in rhabdomeres
I through 6 at 340 to 350 nm and at 480 to 510 nm.
They concluded also that receptor units 7 and 8 were
capable of distinguishing polarized light. We examined
the reflection of sunlight from the surface of clear AI-
synite with a hand-held, polarized plastic sheet and de-
termined that the polarized light is reflected. However,
we did not conduct tests to determine the differences in
fly capture of clear Alsynite panels that reflected polar-
ized light and those that did not.
Of the II colors field evaluated, none was as efficient
as the clear Alysnite (Table I). The reflectances of the
glossy white and light pink were almost identical in the
350 to 370-nm region, with glossy white reflecting < 10%
more than pink. However, white exhibited a 90% re-
flectancy at 450 nm, whereas pink was only ca. 50% at
400 nm. The yellow increased only ca. 2% in reflec-
tancy in the 350 to 420-nm range, whereas red exhibited
FLY VISION AND TRAPPING 1827
no change until - 550 nm (Fig. 5A). Thimijan et al.
(1973) measured the behavioral response of house, sta-
ble. and face flies to different wavelength bands in the
laboratory and determined that maximum numbers re-
sponded to 330-nm stimuli. Muniz and Hecht (1968)
reported that stable flies were attracted to the following
colors in descending attractancy-red, brown, green,
black, yellow, grey, white, and blue. When the spectral
reflectance curves were plotted, the colors that were
most attractive had a reflectance of less than 20% in the
range from 350 to 450 nm. Waldbilling (1968) found
that stable flies mainly responded to colors in the range
from 340 to 500 nm, with a peak at 420 nm. Our field
behavioral studies indicate that surfaces with UV re-
flectancy of ca. 10 to 20% in the range from 360 to 420
nm are attractive to stable flies. These flies exhibit peak
electrophysiological response in the 360-nm region and
again at ca. 490 and 625; however, the region from 360
to 400 nm is the most attractive behavioral stimulus.
During cool weather, the flies may be responding more
to the 625-nm region for body temperature thermore-
gulation (Bushman and Patterson 1981).
Although it was not the goal of this study on the
stable, horn, and face flies to track the information flow
in neural processing areas of the flies' central nervous
systems, Kirschfeld (1972) has studied the structure and
function of the neural elements connecting the eight
rhabdomeres that form the single visual unit, the om-
matidium in the eye of M. domestica. He used opto-
motor studies to demonstrate differences in information
output from rhabdomeres 1 through 6 and 7 and 8, and
concluded that there were differences between the two
receptor groups concerning absolute and spectral sen-
sitivity, sensitivity to polarized light, and contrast trans-
fer.
In our field tests, hom flies were found in abundance
on cattle in pastures, yet fewer than I % were captured
on trap surfaces. In the presence of a host, horn flies
were not attracted to trap surfaces that had color reflec-
tances that spanned the spectrum from UV to red, though
the eye of the horn fly was quite responsive to UV
wavelengths. However, horn and stable flies in a labo-
ratory room can be removed within minutes by using a
point source of white or UV light in a dark room (Pat-
terson and Miller 1982).
The peak visual sensitivities of the stable, horn, and
face flies occur in the UV and blue-green regions, and
there is a correlation between the percent reflectance of
the surface of efficienllraps and visual sensitivity of the
stable fly. Stable flies are attracted to surfaces that have
a 5 to 20% reflectance in the UV range from 350 to 420
nm. This agrees with Waldbilling (1968), who found
that stable flies were attracted to light stimuli of 420
nm. Face flies also exhibit a similar response to surfaces
reflecting in the UV range according to Pickens (J 983).
We hope this study will stimulate others who are de-
veloping better insect traps to consider the visual sen-
sitivity of the insect being trapped, the specific reflectance
of the trap surface, and specific behavior of the insect
in the field.
1828 ENVIRONMENTAL ENTOMOLOGY
Acknowledgment
We thank L. G. Pickens for his helpful suggestions, John
C. Davis for his assistance in collecting and summarizing the
vision data. and Elaine S. Turner for editorial suggestions and
manuscript preparation.
REFERENCES CITED
Agee, H. R. 1973. Spectral sensitivity of the compound eye
of field-collected adult bollworms and tobacco budworms.
Ann. Entomo!. Soc. Am. 66: 613-615.
Ballard, R. C. ]958. Responses of Stomoxys ca/citrans to
radiant energy and the relation to adsorption characteristics
of the eye. Ibid. 5: 449-464.
Bidgood, H. M. 1980. Host location in Stomoxys ca/citrans
L. (the stable fly). Ph.D. dissertation. University of Bir-
mingham. United Kingdom. 152 pp.
Bushman, L. L., and R. S. Patterson. 1981. Assembly. mat-
ing. and thermoregulatory behavior of stable flies under
field conditions. Environ. Entomol. 10: 16-21.
Driggers, D. P. 1971. Field evaluation of blacklight electro-
cutor grid traps for the control of flies associated with poul-
try. Master of Science thesis. University of Florida.
Gainesville. 34 pp.
Gatehouse, A. G., and C. T. Lewis. 1973. Host location
behavior of Stomoxys ca/citrans. Entomo!. Exp. App!. 16:
275-290.
Goldsmith, T. H., and H. R. Fernandez. 1966. Some pho-
tochemical and physiological aspects of visual excitation in
compound eyes. pp. 125-143. In C. G. Bernhard [ed.] .•
Functional organization of the compound eye. Pergamon
Press. Oxford.
Hansen, E. J. 1951. The stable fly and its effect on seashore
recreational areas in New Jersey. J. Econ. Entomol. 44:
482-487.
LaBrecque, G. C., D. W. Meifert, and J. Rye, Jr. 1972.
Experimental control of the stable fly. Stoxomys ca/citrans
(Diptera: Muscidae) by releases of chemosterilized adults.
Can. Entomo!. 104: 885-887.
Kirschfeld, K. 1972. The visual system of Musca: Studies on
optics, structure and function. pp. 61-74. In R. Wehner
[ed.], Information processing in the visual system of ar-
thropods. Springer-Verlag. Berlin.
McCann, G. D., and D. W. Arnett. 1972. Spectral and po-
larization sensitivity of the dipteran visual system. J. Gen.
Physiol. 59: 534-558.
Vol. 12, no. 6
Meifert, D. W., R. S. Patterson, T. Whitfield, G. C.
LaBrecque, and D. E. Weidhaas. 1978. Unique attrac-
tant-toxicant system to control stable fly populations. J.
Econ. Entomo!. 71: 290--292.
Miller, J. A., C. D. Schmidt, C. D. Eschle, Jr. 1979. Mass
rearing of hom flies on a host. U.S. Dep. Agrie. Tech.
Rep. No.8.
Muniz, R., and O. Hecht. ]968. Observations on the dimi-
bution of the stable fly. Stomoxys ca/citrans. on a small
farm and trial on selection of colored surfaces in free air.
Ann. Esc. Nac. Ciene. Bio!' Mexico 17: 225-243.
Parr, H. C. M. 1962. Studies on Stomoxys ca/citrans L. in
Uganda. E. Africa. III. Notes on the life history and be-
haviour. Bull. Entomo!. Res. 53: 437-443.
Patterson, R. S., and O. Juma. 1980. Control, biology and
ecology of Stomoxys flies S. negra Marquart and S. ca/ci-
trans L. (Diptera. Muscidae) on Zanzibar, Tanzania. USAID
Report AIP698-410. 27 pp.
Patterson, R. S., and J. A. Miller. 1982. The sterile insect
technique in integrated pest management programmes for
the control of stable flies and hom flies. In Sterile insect
technique and radiation in insect contro!. Proc. Symp. Neu-
herberg 1981. IAEA, Vienna. 495 pp.
Pickens, L. G. 1983. Face fly response to colors. Entomo!.
Exp. App!. 33: 11-16.
Pickens, L. G., R. W. Miller, and J. J. Grasela. 1977.
Sticky panels as traps for Musca autumnalis. J. Econ. En-
tomo!. 70: 549-552.
Pospisil, J., and J. Zdarek. 1965. On the visual orientation
of the stable fly Stomoxys calcitralls (L.) to colors. Acta
Entomol. Bohemoslor 62: 85-91.
Ruff, J. P. 1979. Trapping effectiveness of several combina-
tions of colors and textures of sticky traps for stable flies,
Stomoxys ca/citrans, Mosq. News 39: 290--292.
Sokal, R. R., and F. J. Rohlf. 1969. Biometry: the principles
and practice of statistics in biological research. W. H. Free-
man & Co .• San Francisco. 776 pp.
Thimijan, R. W., L. G. Pickens, and N. O. Morgan. ]973.
Responses of the house fly. stable fly. and face fly to elec-
tromagnetic radiant energy. J. Ecan. Entoma!. 66: 1269-
1270.
Waldbilling, D. C. 1968. Color vision of the female stable
fly. Stomoxys ca/citrans. Ann. Entomo!. Soc. Am. 61: 789-
791.
Williams, D. F. 1973. Sticky traps for sampling populations
of Stomoxys ca/citrans L. J. Econ. Entomo!. 66: 1279-
1280.