Veterinary Parasitology 91 (2000) 93–105

Factors affecting the larvipositional activity of

Oestrus ovis gravid females (Diptera: Oestridae)
R. Cepeda-Palacios a,∗ , P.J. Scholl b
a Depto. de Zootecnia, Universidad Autónoma de Baja California Sur, Km. 5.5 Carr. al Sur, P.O. Box 19-B,
23080 La Paz, Baja California Sur, Mexico
b Fort Dodge Animal Health, P.O. Box 400, Princeton, NJ 08543-0400, USA

Received 21 January 2000; accepted 13 April 2000

Abstract
Observations of fly strikes or larvipositions (n=68 in 21 days of observation) were carried out in a
herd of goats during the spring in Baja California Sur, Mexico in order to identify the climatic condi-
tions favoring larviposition activity of gravid Oestrus ovis L. flies, as well as to investigate whether
a mixture of some potentially useful compounds was involved in this behavior. Hand-caught, teth-
ered flies (n=43) were either exposed or unexposed to a combination of carbon dioxide, humidity,
1-octen-3-ol, butyric, propionic, acetic acid and acetone released from movable sheep and goat
dummies under open field and cage conditions. Fly strikes occurred at temperatures greater than
20◦ C, but mainly between 25 and 28◦ C and from 116 to 838 W m−2 of solar irradiance. Few or
no strikes were seen under moderate or strong wind, but did occur in a wide range of relative
humidity. The chemicals applied did not improve the capacity of animal dummies to induce the
flies to larviposit, but very irregular behavior was observed. Fourteen larvipositions were made
on the dummies lacking chemical stimuli, so visual ability and movement by the dummies was
very important in stimulation of the flies. Temperature appeared to be the main factor determining
fly activity, but wind and solar irradiance also played important roles. Characteristics of O. ovis
larviposition are discussed. © 2000 Elsevier Science B.V. All rights reserved.
Keywords: Sheep nose bots; Adult; Stimulation; Behavior; Climatic factors; Goats

1. Introduction

Oestrus ovis L., the sheep nose bots, are myiasis-causing dipterans that are widely dis-
tributed among sheep and goat populations in the world (Zumpt, 1965; Touré, 1994). Control
of this parasite has focused mainly on the larval stages and has ignored opportunities to
∗ Corresponding author. Tel.: +52-112-80802; fax: +52-112-80880.

E-mail address: rcepeda@calafia.uabcs.mx (R. Cepeda-Palacios)

0304-4017/00/$ – see front matter © 2000 Elsevier Science B.V. All rights reserved.
PII: S 0 3 0 4 - 4 0 1 7 ( 0 0 ) 0 0 2 6 5 - X
94 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105

focus on the non-parasitic pupal and adult stages. This may be because scant information
is available about the field behavior of O. ovis, and even less about the mechanisms that
control specific activities such as host location and larviposition by gravid females. O. ovis
males and gravid females can be seen on board walls and stones or at rest on buildings, si-
los, tree trunks, and rocks (Porchinskii, 1913; Babcock, 1953; Grunin, 1959), and the flight
radius has also been studied (Semenov et al., 1975). The majority of authors agree that
larvipositing flies are active mainly during sunny, hot days (Cobbett and Mitchell, 1941;
Babcock, 1953; Zumpt, 1965). More accurate information on this behavior is necessary
when the use of attractants for trapping adults is desired. Trapping O. ovis flies might con-
stitute one measure for population monitoring and control given the capacity of females to
produce more than 500 eggs in a relatively short life-span (Cobbett and Mitchell, 1941).
For another oestrid fly, Cephenemyia spp., carbon dioxide was reported to attract gravid
flies when used in Arnaud insect flight traps (Anderson and Olkowski, 1968), and larvipo-
sition was achieved with carbon dioxide combined with 1-octen-3-ol (octenol) and deer
scent using deer models (Anderson, 1989). Recently, the capability of CO2 as an attractant
for the gravid oestrid parasites of reindeer Cephenemyia trompe Modeer and Hypoderma
tarandi L., was demonstrated (Anderson and Nilssen, 1996a,b). These studies also demon-
strated that climatic factors, hour of the day, and physiological age of the flies significantly
influenced the rate of trapped flies. Our objectives were, therefore, to identify the climatic
conditions favoring larviposition activity of O. ovis gravid females and to determine if a
mixture of some potentially useful compounds could stimulate them to larviposit.

2. Materials and methods

2.1. Description of the study site

Observations were carried out during the spring of 1999 on a ranch located at 26◦ latitude
N, 111◦ 500 longitude W, and altitude 200 m, in Baja California Sur, Mexico. Locally, spring
is the season with the highest O. ovis fly activity (Cepeda et al., 1993). The climate is warm
and dry (200 mm average annual rainfall) with the major precipitation events occurring at
the end of the summer and minor precipitation in the winter. A herd of about 350 adult goats
grazed during the morning and part of the afternoon within a radius of ca. 5 km from an
open, 12 m×10 m corral where they were gathered at night. The pasture consisted of a hilly,
arbocrasicaulescent shrubland where dominant plant species included Prosopis palmeri,
Cercidium microphyllum, Opuntia cholla, Pachycereus pringlei and Fouquieria diguetti
(Aguirre, 1975). In the afternoon the breeding stock usually returned from the pasture in
groups of varying numbers and drank water in a well located ca. 60 m from the corral. As
occurred in the rest of the study area, the distance between the corral and the well was
covered with shrubs, logs, and rocks that might have harbored infective flies.

2.2. Observations of fly strikes

Fly strikes were observed during the morning (generally from 07:00 to 09:00 hours or
later if goats were available) and in the afternoon (mostly from 15:00 hours to dusk) when
 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105 95

Table 1
Description of the wind scale used to classify the wind conditions when Oestrus ovis strikes occurred
Wind categorya Speed (km h−1 ) Characteristics

1 1.6 Calm, no movement in leaves of trees

2 1.6–5.0 Sustained light air, slight motion in leaves
3 3.0–9.0 Light air with gusts of light breeze
4 6.4–11.0 Sustained light breeze making leaves rustle
5 13.0–19.0 Gentle breeze, constant movement in leaves and twigs
6 16.0–25.0 Gentle breeze with gusts of moderate breeze
7 21.0–29.0 Sustained moderate breeze, small branches are moved
a Adapted from the Beaufort Wind Scale described by Byers (1974).

the goats were housed in the corral. Striking flies appeared suddenly (especially when the
herd walked to enter the corral), usually larviposited on several goats, having short time
intervals between larvipositions and then disappeared outside the corral. During the periods
of rest, the flies were seen on the ground among goats or on goats, showing grooming
and alert behavior. A fly strike was noticed when goats began to sneeze as a defensive
reaction and flies were seen among them. Immediately after this happened, the number
of goats present in the corral was recorded along with the air temperature, peak solar
irradiance (using an Eppley radiometer, New Port, RI), relative humidity (using a dry-wet
bulb sling psychrometer) and wind velocity using a seven-category scale (Table 1) because
an anemometer was not available. Even if no fly strikes were observed, these variables
were recorded at irregular intervals. Measuring total solar irradiance was preferred over
measuring light intensity because only 46% of solar radiation is in the visible wavelength
band while 49% is in the infrared and the rest is in the ultraviolet (IEA, 1987). Thus, solar
irradiance provides both heat and light to the exposed fly. It has been reported, for example,
that when bot flies have been exposed to direct solar light, the temperature threshold for
flight is modified (Breyev, 1956 cited by Anderson and Nilssen, 1996a). When possible,
larvipositing flies were hand-caught and set in cardboard cages for further trials. For this
purpose, a skilled person waited for a larvipositing fly to perch on the ground or on a quiet
goat, then he slowly approached the fly to catch it with a rapid movement of hand. Relatively
immobile goats were a necessary condition to catch the flies. A total of 68 fly strikes were
recorded during a period of 21 days of observation.

2.3. Visual and chemical stimulants of fly larviposition

Two dummy models were used, one a Suffolk breed-type sheep made of synthetic resin
and the other a taxidermically-prepared, Anglo-Nubian breed-type goat. Both models were
fitted to release CO2 gas, humidity, octenol (Aldrich Chemical Co., Milwaukee, WI), the
volatile fatty acids (VFAs) butyric (Merck-Schuchardt, Germany) propionic, acetic, as well
as acetone (Productos Quı́micos Monterrey, Monterrey, Mexico), with olive oil added as
a vehicle. Octenol has been used for attracting gravid Cephenemyia spp. flies as well as
tsetse flies (Hall, 1995). Volatile fatty acids and CO2 are products of the ruminal microbial
activity on feed components and of host metabolism, respectively, whereas acetone may be
found in the breath and urine of stressed ruminants under high energy output (Van Soest,
96 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105

1994), the condition known as ketosis. All trials were done under direct solar irradiance
(>407 W m−2 ) and at air temperatures above 23◦ C. The models were fixed with metal
spring devices that allowed them to move so as to imitate live animal head movements.
Carbon dioxide was released (4 l/min) from a hose stuck on the muzzle of the models that
was connected to a gauged container. The volatile chemicals were separately prepared as
10% (v/v) of the compound in olive oil. At the precise moment of initiation of chemical
stimulation of larviposition, 2 ml of each volatile chemical preparation was applied into a
sponge which was fixed under the chin of the dummy. Humidity was applied by forcing
carbon dioxide to flow through a small warm plastic chamber containing a sponge saturated
with distilled water. The combination of volatile chemicals was renewed every 15 min if a
trial lasted more than this period of time.

2.4. Stimulation of larviposition

After collecting gravid flies, several modalities for testing the stimulating effects of
different chemical compounds were attempted. In the beginning, freshly caught wild flies
were transported to the laboratory at La Paz, B.C.S., 300 km from the collection site to be
tested the morning after, but rarely did the flies remain alive more than 12 h after being
caught. After this period, few live flies were able to fly appropriately and the remainder
never adapted to captivity. Surviving flies failed to fly toward the models whether with or
without the chemical stimulants, and their activity consisted only of escape attempts. Next
we tried open field operations at the ranch with tethered flies (Anderson, 1975; Nilssen
and Anderson, 1995) in which the flies, when necessary, were first immobilized with CO2
in order to attach a 1 m long polyester thread onto the scutum. The models were fixed to
fence poles, then, testing the stimulating capacity of the models alone (to avoid any further
residual chemical stimulation, firstly using the caprine, then the ovine model) was tried
and finally, the mixed candidate stimulants were applied on the models. Some attempts to
induce larviposition of tethered flies on live goats were also made. The fly being tested was
allowed to fly freely around the dummy within the range of the thread. When a larviposition
occurred, the time elapsed from the beginning of the test, the number of larvae deposited and
the side of the nostril in which the larvae were found, was recorded. Because some poorly
attached flies escaped and because sometimes it was difficult to operate in strong wind, the
final attempt to testing the candidate compounds was inside a walk-in cage (1.22 m×2.45 m
base, 1.8 m height) covered with transparent polyethylene to allow illumination from the
sunlight. Solar irradiance and air temperature inside the cage were continuously monitored.
Of 69 available flies, only 43 had satisfactory flight capacity and were used in 29 sessions
that lasted about 38 h in total. Some selected sessions were videotaped and later analyzed.

2.5. Data analysis

Air temperature, solar irradiance, relative humidity and wind data were plotted against
the frequency of observed strikes. Descriptive statistics of these variables when the strikes
occurred were calculated. Because of the great variability and unpredictable behavior and
life-span observed with the tethered flies, no experimental design could be used. Results of
stimulated larvipositions are only descriptively presented.
 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105 97

3. Results

3.1. Temperature

Within a range of 9.0–36◦ C (Fig. 1A), fly strikes were observed only when temperatures
exceeded 20◦ C. The major peak of strikes was seen between 25 and 28◦ C (the maximum
number of strikes occurred at 26◦ C), but a few strikes occurred at 34 and 35◦ C. During

Fig. 1. Influence of air temperature (A) and solar irradiance (B) on the frequency of Oestrus ovis fly strikes observed
on goats during spring in Baja California Sur, Mexico.
98 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105

spring, air temperatures were seldom below 9◦ C or over 35◦ C, and on cloudy, cold days in
which the diurnal temperature did not reach 20◦ C, no flies were seen around the goats.

3.2. Solar irradiance

Strikes occurred in a range from 116 to 838 W m−2 (Fig. 1B) which corresponded to
irradiance prior to sunset and strong sunlight, respectively. This meant that the gravid flies
were able to infect goats until late in the day and on cloudy days when other climatic
factors (i.e. temperature and wind) permitted larviposition. Two major peaks of activity
were observed, one at 558 W m−2 and the other at 686 W m−2 .

3.3. Wind speed

Flies were able to larviposit under conditions of calm to moderate wind (Fig. 2A). How-
ever, in stronger wind, strikes were absent. The number of fly strikes during gusts of light
breeze was rather low because very few observations were obtained during this condition.

3.4. Relative humidity (RH)

Larviposition strikes occurred in a very wide range of relative humidities (Fig. 2B).
Within a tested range of 44–100% RH, the flies actively larviposited between 51 and 96%
RH, with higher peaks between 65 and 85% RH.

3.5. Time of day

Although observations were totally dependent on the presence of goats at the place of
study, it is interesting to note that most strikes occurred between 14:30 and 19:30 hours,
corresponding to the warmest time of day in spring prior to dusk (Fig. 3A), only when the
goats had returned from grazing. Few strikes were seen in the morning before the goats left
the pen for grazing. No O. ovis flies were seen around the pen or ranch house when no goats
were present.

3.6. Number of goats in the pen

A noteworthy observation was the number of observed strikes seen relative to the num-
ber of goats that gathered together following grazing (Fig. 3B). When the number of goats
gathered was low (i.e. <45), no strikes were seen. The probability of observed strikes was ob-
served to be higher when more goats (80 or more) came together from the field. Descriptive
statistics for records of climatic factors when fly strikes occurred are summarized in Table 2.

3.7. Stimulation of fly larviposition

The candidate stimulant combination applied did not improve the capacity of the mod-
els to induce females to larviposit (Table 3). In total, seven different flies responded by
 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105 99

Fig. 2. Influence of wind (A), represented in seven categories from 1, calm (1.6 km h−1 ) to 7, moderate breeze
(21–29 km h−1 ) and relative humidity (B) on the frequency of Oestrus ovis fly strikes observed on goats during
the spring in Baja California Sur, Mexico.

larvipositing from one to six times (four flies larviposited on the goat model and three
flies did so on both the sheep and goat models). Fourteen larvipositions occurred on the
models with no chemical stimulation, only three on the same models with chemicals and
two larvipositions by two flies were achieved on live goats in the open field. Two larvipo-
sitions on the models (obtained from two flies, using the chemical combination) could not
be repeated by the same flies with or without the chemicals. Some flies readily larviposited
several times on the models. For example, a tethered fly larviposited four times on OD and
100 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105

Fig. 3. Frequency of Oestrus ovis fly strikes occurring at different times of the day (A) and different number of
goats present in pen (B) when the strikes occurred.

only once on CD in the open field; another fly larviposited once on MC+CD, once on CD
and three times on OD. For this reason, the number of flies that larviposited at least once
in each modality is also presented in Table 3. On an average, 10.4±6.8 larvae per strike
were deposited (range 3–24) with the same proportion in the right or left nostril. A strike
consisted of a sticky package of closely joined larvae that quickly began to spread on the
muzzle following larviposition, probably looking for heat and/or humidity. The time of
reaction (interval from onset of the release of the tethered fly to initial larviposition) was
 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105 101

Table 2
Descriptive statistics of climatic variables when Oestrus ovis strikes occurred
Variable n Mean s

Temperature (◦ C) 68 27.1 3.8

Wind speeda 68 3.4 1.5
Solar irradiance (W m−2 ) 52 522.0 188.0
Relative humidity (%) 52 74.0 13.0
Number of goats in pen 59 115.0 53.0
a Wind category 3 corresponds to approximately 3–9 km h−1 .

13.4±9.7 min (range 2–30 min); in those flies having subsequent larvipositions, the time
interval was 7.3±4.8 min (range 2–15 min). Because movement was manually induced in
the models, sometimes the models remained motionless for short periods of time until
movement resumed. Movement of the models was necessary in order to attract attention of
the flies during stimulation. Only one larviposition was accomplished with no movement
by the goat model and none in the sheep model. Some very active flies with extremely
irregular flight patterns larviposited in just one strike without stopping in front of the nose
of the models. Other flies showed momentary stationary flight patterns or hovered in front
of the model muzzle, followed by a small movement forward of the fly to eject the larvae
on the host’s nostril. The estimated distance between the flying female and the target was
from 1 to 3 cm.

Table 3
Stimulation of larvipositions of Oestrus ovis flies, carried out under different modalities of stimulus application
Modality n (utile)a Temperatureb (◦ C) SIb (W m−2 ) Stimulic Total LVPd Fliese
Walk-in cage, free flies 10 (5) 29 407 LG 0 0
MC 0 0
CD 0 0
Open field, tethered flies 32 (19) 27 523 LG 2 2
MC+CD 1 1
MC+OD 1 1
CD 1 1
OD 4 1
Walk-in cage, tethered flies 27 (19) 30 465 MC+CD 1 1
MC+OD 0 0
CD 6 4
OD 3 1
a Number of flies used in each modality and number of flies with adequate flight in parenthesis.
b Mean temperature and solar irradiance (SI) under which stimulations were carried out.
c LG, Live goat; MC, mixture of chemicals and CO ; CD, use of the goat dummy; OD, use of the sheep dummy.
2
d Total LVP is total number of larvipositions obtained is each modality.
e Flies stands for number of flies that larviposited at least once in each modality.
102 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105

4. Discussion

Among the climatic factors influencing oestrid fly activity, temperature, light intensity,
and wind are recognized as the most important. Anderson and Nilssen (1996a) determined
critical values of temperature, light intensity and wind velocity for C. trompe and H. tarandi
fly activity in northern Norway; no fly activity was observed during periods of rain or snow.
In the case of O. ovis, it has been reported that temperature is the principal factor determining
fly activity. Porchinskii (1913) observed that the flies were active only during the heat of
the day and decreasing temperatures were associated with less activity for both female and
male flies, even on days that were windy, cool or rainy. In a brief report, Babcock (1953)
mentioned that the minimum range for O. ovis fly activity was 16–18◦ C and the optimum
was 27◦ C in Texas. In our study, 20◦ C was the minimum temperature of fly activity while
optimum range appeared to be 26–28◦ C. Adaptive phenomena in fly populations may result
in different temperatures needed for fly activity. In Baja California Sur, for example, diurnal
temperatures in springtime vary from 9 to 35◦ C and, during the year, freezing levels are
rarely reached. Depending on the species, a critical temperature is necessary for flight
activation (Mordue et al., 1980), but individual differences are also common. With O. ovis,
the cold, dormant flies became alert between 11 and 17◦ C and began to follow mobile
objects without flying from 12 to 18◦ C, but the capacity for strong, sustained flight was
achieved only between 16 and 26◦ C in different individuals (unpublished observations).
Because there was no rainfall during the study, either wind velocity or solar irradiance
may be the second most important factor (after temperature) in determining the larviposition
behavior of O. ovis flies. Frequently in the morning, a cold humid wind, low clouds and fog
coming from the Pacific Ocean prevented fly activity until 09:00 or 11:00 hours, even if some
goats were still in the pen. The particular role played by these two factors in influencing
fly activity is interesting. Anderson and Nilssen (1996a) could not trap or observe either
C. trompe or H. tarandi at wind velocities above 8 m/s. When we worked with tethered
flies in the open field, the flies quit flying when gusts of strong wind occurred; the flies
either attempted to escape or they rested on any object and adopted the typical position
of a dormant fly. Moreover, strong winds might have interfered with the precise flight
needed for accurate larviposition. Solar irradiance, and therefore light intensity, may be
important for host-seeking and larviposition activities in the female as a limiting factor for
performing these tasks. The active fly is able to orient the longitudinal body axis toward
gross moving objects as a defensive or normal reaction. We have carried out simple field
tests (unpublished) in which mobile objects (e.g. a person walking) passed in front of alert
flies. This response appeared to be solar irradiance dependent but highly variable among
individual flies which showed maximum visual range of 6–7 m, under solar irradiances
of 140–1160 W m−2 . Except during very humid weather, relative humidity did not seem
to limit fly activity. The flies were able to larviposit under a very wide range of relative
humidity values.
The larviposition act itself may be regarded as a learned or innate complex process in
which behavioral, olfactory, visual and/or environmental stimuli may be involved. In this
study, we found that visual stimulus alone may be sufficient to elicit this response in some
flies and that the chemical mixture used did not modify larvipositional behavior in the flies.
Cobbett (1948), on the other hand, concluded that tar or tar mixed with oils, fish oils and
 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105 103

other repellents applied to sheep noses were not effective in prevention of larviposition.
Although fly attraction and stimulation must be separated in studies, these results agree
in part with those obtained by Anderson (personal communication) in that CO2 released
from tent-like Arnaud flight traps was unable to attract host-seeking O. ovis. Based on
examples reported in the literature, speculations on why flies larviposit on models can be
made. Touré (1994) pointed out that gravid females showed tropism toward the nostrils of
small ruminants. Nicoli et al. (1962) discussed human ophthalmomyiasis caused by O. ovis
and suggested that, when lacking normal sheep or goat hosts, the flies deposited larvae in
human eyes because of the ‘configuration of the human face’. Zumpt (1965) discarded the
idea that sheep odors on shepherds were responsible for attracting gravid flies to larviposit
on them because the flies also larviposited on persons with no connection with normal
host species. The lack of consistency of the responding flies to larviposit repeatedly on any
target may be mainly attributed to their wild origin. As expected, captive flies showed a
very modified behavior compared to free, larvipositing flies. Tethered and caged flies were
always actively engaged in escaping, even when live potential hosts were present. On the
other hand, variable degrees of fly inconsistency may be observed even in highly controlled
experiments (Dethier, 1974). Slightly more flies larviposited on models without chemicals
in the walk-in cage. This was due to the order in which stimuli were applied to stimulate
the flies (models without chemicals first), the same flies had lost flight ability when the
chemical mixture was subsequently applied.
Color of the target, age of the fly and previous larviposition experience may affect the
larviposition response. Meleney et al. (1962) reported less severe O. ovis infections in black
sheep. This possible color preference, together with the fact that the captured flies had
been larvipositing on goats only, might have influenced the lack of response in some flies
to larviposit on the black-faced sheep model. However, Murguı́a et al. (2000), based on a
seroprevalence study, reported that sheep with dark noses showed a higher probability of
being infected, compared to sheep having light-colored noses. Unlike C. trompe and H.
tarandi (Anderson and Nilssen, 1996b), assessing the relative age of wild, larvipositing O.
ovis flies using the remains of abdominal fat body is difficult because it was totally spent
during the activities prior to larviposition. Dissection of the flies showed that only 8% of
them had dropped their larval reserves, the rest had 25 to almost 100% of their original
component of larvae remaining in utero. Movement of potential hosts, such as walking in
herds and/or stimulation such as movement of the legs and heads, may be a very important
cue in attracting larvipositing females. These same responses have been observed in H.
tarandi (Anderson and Nilssen, 1996a), but this still is a largely open subject for future
research (Hall, 1995). On several occasions in the present work, we were able to catch
more flies by making the herd of goats walk through the shrubs and big stones located
between the water well and the pen. Based on these observations, we suggest that after
the gravid fly–host encounter, the former larviposits (with favorable weather) continuously
and remains around or moves with the herd until its larval reserve is depleted, probably
within 1 or 2 days at most. Field observations by the goat keepers indicate that the flies
frequently travel on the rumps or horns of grazing goats. Finally, it still is not known whether
long-distance or short-distance attractants or stimulant compounds and/or visual cues are
involved in the processes of host seeking or larviposition. Research on live animal body
secretions and visual stimuli which has been done for other myiasis-producing genera such
104 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105

as Cochliomyia spp. and Lucilia spp. blowflies (Green and Warnes, 1992; Wall et al., 1992;
Wall and Smith, 1996; Morris et al., 1997), is lacking for oestrid species.
Evidence here presented indicated that temperature was the chief climatic factor involved
in the larvipositional activity of O. ovis females, followed by solar irradiance and wind
speed. Under local conditions, 20◦ C appeared to be the critical temperature for activation
of larvipositional activity. Movement by the potential host also played an important role in
provoking larviposition behavior.

Acknowledgements

We gratefully acknowledge the assistance of Loreto Andrade M. and Stephanie Frugère

during the field work; and of Salomon Murillo M. and Lorenzo Murillo M. for long, il-
lustrative talks. The authors are also especially grateful to Dr. M.F. Chaudhury from the
Screwworm Research Lab., Tuxtla Gutiérrez, Chiapas, Mexico and Dr. J.R. Anderson for
valuable advice.

References

Aguirre, M.R., 1975. Coeficientes de Agostadero de la República Mexicana. Estado de Baja California Sur. S.A.G.,
México, D.F., 120 pp.
Anderson, J.R., 1975. The behavior of nose bot flies (Cephenemyia apicata and C. jellisoni) when attacking
black-tailed deer (Odoicoileus hemionus columbianus) and the resulting reactions of the deer. Can. J. Zool. 53,
977–992.
Anderson, J.R., 1989. Use of deer models to study larviposition by wild nasopharyngeal bot flies (Diptera:
Oestridae). J. Med. Entomol. 26, 234–236.
Anderson, J.R., Nilssen, A.C., 1996a. Trapping oestrid parasites of reindeer: the response of Cephenemyia trompe
and Hypoderma tarandi to baited traps. Med. Vet. Entomol. 10, 337–346.
Anderson, J.R., Nilssen, A.C., 1996b. Trapping oestrid parasites of reindeer: the relative age, fat body content and
gonotrophic conditions of Cephenemyia trompe and Hypoderma tarandi females caught in baited traps. Med.
Vet. Entomol. 10, 347–353.
Anderson, J.R., Olkowski, W., 1968. Carbon dioxide as an attractant for host-seeking Cephenemyia females
(Diptera: Oestridae). Nature 220, 190–191.
Babcock, O.G., 1953. Notes on the biology of Oestrus ovis L. Bull. E-871, USDA-ARS. Texas Agricultural Station.
Byers, R.H., 1974. General Meteorology, 4th Edition. McGraw-Hill, New York, 260 pp.
Cepeda, R., Greene, E., Loya, J.G., Espinoza, J.L., 1993. Incidencia de Oestrus ovis en rebaños de ovinos y
caprinos en Baja California Sur. Rev. Inv. Cient. Cs. Agrop. 4, 29–36.
Cobbett, N.G., 1948. The sheep head grub and methods of control. Bureau of Animal Industry, USDA, Mimeo.
Cobbett, N.G., Mitchell, W.C., 1941. Further observations on the life cycle and incidence of the sheep bot, Oestrus
ovis, in New Mexico and Texas. Am. J. Vet. Res. 2, 358–366.
Dethier, V.G., 1974. Sensory input and the inconstant fly. In: Browne, L.B. (Ed.), Experimental Analysis of Insect
Behaviour. Springer, Berlin, pp. 21–31.
Green, C.H., Warnes, M.L., 1992. Responses of female New World screwworm flies, Cochliomyia hominivorax,
to colored targets in the laboratory. Med. Vet. Entomol. 6, 103–109.
Grunin, K.Y., 1959. Congregations of bot-fly males on the highest points in locality and the cause. Zoologicheskii
Zhurnal 38, 1683–1688 (Translated from Russian).
Hall, M.J.R., 1995. Review: trapping the flies that cause myiasis: their responses to host-stimuli. Ann. Trop. Med.
Parasitol. 89, 333–357.
IEA (International Energy Agency), 1987. Renewable Sources of Energy. OECD Publications, Paris, 334 pp.
 R. Cepeda-Palacios, P.J. Scholl / Veterinary Parasitology 91 (2000) 93–105 105

Meleney, W.P., Cobbett, N.G., Peterson, H.O., 1962. The natural occurrence of Oestrus ovis in sheep from the
Southwestern United States. Am. J. Vet. Res. 23, 1246–1251.
Mordue, W., Goldsworthy, G.J., Brady, J., Blaney, W.M., 1980. Insect Physiology. Wiley, New York, 108 pp.
Morris, M.C., Joyce, M.A., Heath, A.C.G., Rabel, B., Delisle, G.W., 1997. The responses of Lucilia cuprina to
odours from sheep, offal and bacterial cultures. Med. Vet. Entomol. 11, 58–64.
Murguı́a, M., Rodrı́guez, J.C., Torres, F.J., Segura, J.C., 2000. Detection of Oestrus ovis and associated risk factors
in sheep from the central region of Yucatan. Mexico Vet. Parasitol. 88, 73–78.
Nicoli, R.M., Lanfranchi, J., Jarry, D., 1962. Les myiases humaines à Oestridae (Oestrasis) (Diptera: Brachycera).
Bull. Soc. Entomol. France 67, 108–113.
Nilssen, A.C., Anderson, J.R., 1995. Flight capacity of the reindeer warble fly, Hypoderma tarandi (L.), and the
reindeer nose bot fly, Cephenemyia trompe (Modeer) (Diptera: Oestridae). Can. J. Zool. 73, 1228–1238.
Porchinskii, I.A., 1913. The sheep bot (Oestrus ovis L.): its biology, characteristics, methods of control and relation
to man. Trudy Biuro PO Entomologii 10, 1–65 (Translated from Russian).
Semenov, P.V., Gomoyunova, N.P., Tarasenko, N.N., 1975. The flight radius of Oestrus ovis. Veterinarya 8, 58–59.
Touré, S.M., 1994. Les myiases d’importance économique. Rev. Sci. Tech. Off. Int. Epiz. 13, 1053–1073.
Van Soest, P.J., 1994. Nutritional Ecology of the Ruminant, 2nd Edition. Cornell University Press, Ithaca, 476 pp.
Wall, R., Smith, K.E., 1996. Colour discrimination by the sheep blowfly Lucilia sericata. Med. Vet. Entomol. 10,
235–240.
Wall, R., Green, C.H., French, N., Morgan, K.L., 1992. Development of an attractive target for the sheep blowfly
Lucilia sericata. Med. Vet. Entomol. 6, 67–74.
Zumpt, F., 1965. Myiasis in Man and Animals in the Old World. Butterworths, London, 267 pp.