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FLIES BREEDING IN WASTES OF
CATTLE & OTHER LIVESTOCK
Musca autumnalis,
Musca vetustissima, Haematobia spp. -- Muscidae (Contacts) Please CLICK on underlined categories to view further details] [Refer also to Related
Research #1, #2 ] The exophilic flies
are those that persist in nature in the absence of humans, but whose
populations can increase as a result of certain human activities such as provision
of greater breeding habitat. They
include several species in the genera Calliphora,
Hippelates, Musca, Muscina, Phaenicia,
and Stomoxys. Some success has been recorded with the use
of natural enemies against the calliphorid species in California and Hawaii,
but attempts elsewhere have not been effective (Bay et al. 1976). The braconid parasitoid Alysia ridibunda Say, indigenous to the United States, was
released into an area of Texas new to its range and successfully parasitized
the blowflies Phaenicia sericata (Meigen) and a Sarcophaga species. However, the parasitoid did not maintain
control and became rare within a couple of years (Lindquist 1940). The gregarious parasitoid
Tachinaephagus zealandicus may have
considerable potential for biological control of exophilic flies (Olton &
Legner 1974, 1975 ). The range of habitats utilized by this
natural enemy is considered unparelleled by any other fly parasitoid. But this genus has not been given much
attention. One species, Tachinaephagus stomoxcida Subba-Rao provides
overall permanent reductions of Stomoxys
in Mauritius (Greathead & Monty 1982).
The complex of
problems that confront field programs in biological control of exophilic
flies has clearly had a dampening effect on research in this area. The unforeseen problems associated with
attempts to biological control the eye gnat, Hippelates collusor
(Townsend), in California exemplify those problems. In the early 1960's a concerted effort was launched to control
this eye gnat with the use of both indigenous and exotic parasitoids in
orchards and date palm groves of southern California. About a dozen species and strains were
evaluated for several years. Some of
the exotics established, but eye gnat reductions were obvious only where
cultivation practices were curtailed (Legner et al. 1966, Legner 1970b). Cultivation of the orchards buried the
larvae and pupae of the eye gnat below the search zone of the parasitoids and
cultivation also removed vegetation that offered the parastioids protection
and possibly nutrients (Legner & Olton 1969, Legner & Bay 1970). Buried eye gnats emerged from several
centimeters below the soil surface and thus continued to pose a serious
problem (Bay et al. 1976). Tabanidae or
horseflies, although widespread and on occasion serious pests and vectors of
disease of livestock, have not received much attention. Only one successful inundative release of
the egg parasitoid Phanurus emersoni Girault has been
recorded (Parman 1928). Apparently
this effort was precipitated by a severe outbreak of anthrax at the time and
since this disease diminished and other control tactics were available,
interest in their biological control has not continued. Flies associated
with cattle droppings, symbovine flies (Povolny 1971), have received the most
attention for biological control since the 1970's. The primary targets for control have been the bush fly, Musca vetustissima Walker, the
hornfly, Haematobia irritans (L.), and the facefly,
Musca autumnalis DeGeer (Wallace & Tyndale-Biscoe 1983,
Ridsdill-Smith et al. 1986, Ridsdill-Smith & Hayles 1987). Scarab beetles have
been the principal emphasis for biological control of pasture breeding
symbovine flies since Albert Koebele first imported coprophages and fly
predators from Europe to Hawaii in 1909 (Anderson & Loomis 1978;
Bornemissza 1976; Ferrar 1975;
Waterhouse 1974; Legner 1986). The
largest effort took place in Australia where pasture improvement benefits
were also desired (Bornemissza 1960 1976; Ferrar 1975). However, widespread significant fly
reductions have not been reported (Legner 1978a 1986; Macqueen 1975). Field and
laboratory studies have shown that the survival of symbovine flies can be
experimentally reduced by dung shredding, scattering and burying activities
of scarab beetles (Blume et al. 1973; Bornemissza 1970;
Moon et al. 1980; Hughes et
al. 1978; Ridsdill-Smith
1981; Ridsdill-Smith et al. 1977; Wallace & Tyndale-Biscoe
1983). Macqueen (1975) and Hughes et al. (1978) reviewed several cases in the field where bush
fly, Musca vetustissima Walker reductions
may have resulted from the activities of scarab beetles; and Ridsdill-Smith
& Mathiessen (1984) gave experimental evidence for some reduction by
endemic and imported scarab beetles.
However, the amount of control achieved was generally low. Immigration of bush flies from outside the
experimental area often confounded the results. The only known
biological control reduction of symbovine flies of noticeable magnitude was
reported from Fiji involving a single predator, Hister chinensis
Quensel, that originally had been intended for other dipterous species
(Bornemissza 1968). A minor success
apparently occurred in Hawaii, which involved both dung-burying scarab and
predatory beetles (Legner 1978b). Scarab beetle field
population densities are often high enough to cause significant dung removal
and pasture improvement (Fincher 1981; Fincher et al. 1981;
Kessler 1983; Waterhouse 1974).
However, whether significant symbovine fly reductions are also
achieved is not certain (Legner 1978a 1986; Macqueen 1975). Experimental
Observations
Haematobia irritans (L.) breeding in flood irrigated pastures of
the lower Colorado Desert of southeastern California continues to remain
unacceptably high during warm seasons (>1,000 adult flies per bovine head)
despite the presence of moderately abundant populations of Onthophagus gazella F. This study suggests that densities of >
40-70 adult beetles per dung pad and giving pronounced dung shredding
activity, caused fly mortality of 38-56%.
The continued high abundance of adult horn flies on cattle suggests
that at >50% mortality, the pasture environment still produces sufficient
flies to saturate cattle, although emigration might be reduced. Additional species of scarabs may be
necessary to increase fly mortality.
However, the dung drying activity of existing O. gazella
significantly could interfere with resident staphylinid beetle breeding,
which was significantly lower in pastures where O. gazella
reached densities of 40 per dung pad.
Scarab beetle activity might also impede the introduction of superior
predatory species for biological control. Reasons for the
above conclusions stemmed from observations in the Coachella Valley of
southeastern California, where established populations of Onthophagus gazella F. seem generally
ineffective in reducing adult populations of H. irritans
in irrigated pastures. In the 1970's
the scarab was imported from Hawaii, and establishment quickly followed
(Legner 1986). The species remains
firmly established throughout the Coachella Valley. and dung scattering and
burying by adult beetles in autumn usually begins within an hour of
deposition when pastures are under regular 21-day irrigation. Scarab beetles that remained dormant in
the sandy loam soils, in some cases for six months during irrigation-free
periods in this largely rainfall-free area, become highly active within a
week to 18 days, following renewed irrigation and cattle stocking. Cattle on these
pastures are often stocked at densities exceeding 25 per ha. and left to
graze for 12-14 days. The amount of
dung that is shredded, scattered and buried daily by the 1-cm long beetles is
enormous. By early autumn, beetle
density generally exceeds 40 per fresh dropping, a density in the range where
fly control can be expected in another species, Musca vetustissima
Walker (Wallace & Tyndale-Biscoe 1983).
Ranchers generally have been pleased
with the manner in which the cattle dung is incorporated into the soil, even
though hornfly control seems lacking.
During warm seasons the cattle sustain continuously high densities of
this fly, usually exceeding 1,000 per head in autumn. These densities do not appear very
different from those attained in pastures where scarab beetles are low or
absent. The apparent lack of adult
horn fly control is not understood, especially as beetle densities are
sufficient for fly larval control to begin to take effect. Cattle grazing in
this area was gradually replaced by horse breeding in connection with the
equestrian sport polo, so that by 1988 only about 10% of the former irrigated
pastures were devoted to cattle grazing.
The overall abundance of O.
gazella declined
proportionally, and the beetle population survived in a few isolated
pastures. A unique opportunity to
quantify differences between pastures that sustained scarab beetles and those
in which they were absent or greatly reduced by delayed recolonization,
presented itself in 1989 when some previously abandoned fields were
reconstituted and stocked with cattle. Experiments to
quantify field breeding densities were made in autumn 1989, a time of year
for maximum horn fly and O. gazella abundance in the Lower
Colorado Desert area of southeastern California. Random samples were taken of dung pads shredded by established O. gazella populations where >40 adult beetles attacked a
single pad. These were compared to
unshredded samples from control pastures in which O. gazella
populations had been reduced to <2 per pad through several continuous
years of fallowing. Studies in
previous years had shown that the controls were suitable for maintaining
large populations of O. gazella (Legner 1986 Legner
& Warkentin, unpub. data). There were four
Bermuda grass pastures with established O.
gazella adults and larvae
and four control pastures in the lower Coachella Valley near the towns of
Coachella and Mecca. These pastures,
with a sandy loam soil, ranged from 3-8 ha. in size, and control pastures
were separated from those with long-established O. gazella
pastures by >10 km. Herds were of
mixed breeds, and were stocked at densities of 25-45 per ha., with
supplemental feeding of concentrates.
Samples were taken
over a period of four days from each of four pastures of both types beginning
on September 28, October 26, and November 23, 1989. Twenty fresh cow pads of ca. 1,495 cc (SD 374 cc), were first
marked and then collected after 120 h of exposure to the pasture. Blume et
al. (1970) have shown that
predators and competitors reach a pad within 48 h, and cause most horn fly
mortality within five days by desiccation of the dung. The sampling method
was that of Roth et al. (1983), consisting of
shovel collections both of the manure and the topsoil 5 to 6 cm from the edge
of the pad and 30 cm deep. The manure
and topsoil were sealed in plastic bags and brought to the laboratory. Half the number of samples (10) were
manually broken to separate adult staphylinid predators and O. gazella and to obtain an estimate of their numbers. The other half were placed intact into
emergence sleeve cages in a greenhouse, incubated at 26-29EC, 55-60%
RH and 14:10-h L:D photoperiod for the emergence of adult horn flies. Beetles in the second incubated set were
removed as they left the dung within 6 h of being caged. Adult horn fly
densities on cattle were assessed with 6X binoculars. The average number and oven-dry weights of
horn fly adults emerging per pad from dung collected in both kinds of
pastures was calculated. The number
of adult predatory staphylinids were counted in each pad. Statistical Analyses--Data were transformed to sqr-rt(X + 0.5) and
analyzed for significant differences using an ANOVA F-test (Steel &
Torrie, 1980). Significant
differences were tested at P<0.05. Horn fly adults
were produced from all pastures but significantly lower numbers were from
fields where O. gazella beetles were active
(Legner & Warkentin 1991).
Estimated reductions ranged from 38-56%. There was also a trend for smaller flies to be collected from
pads producing the highest horn fly numbers, based on oven-dry weight data,
bivariate correlation analysis giving a highly significant coefficient of
-0.669, 22 df, P<0.05. Although O. gazella
was either absent or at densities averaging <1 per pad at the beginning of
the sample period, there was a trend toward higher beetle densities on
succeeding sample dates, none of which exceeded 5 per pad (Legner &
Warkentin 1991). Colonization of the
control fields continued in the spring and summer of 1990 so that by July
12th, 1990 the scarab population approximated that observed in the long-established
fields of autumn 1989. This provides
evidence for the suitability of control pastures to sustain equal scarab
densities. Adult hornfly
populations on cattle during the three months sample period remained high
(>1,000/head by 4 PM) in both kinds of pastures. It is unlikely that the large number of horn flies on the
cattle in pastures containing high population densities of O. gazella was due to the immigration of flies from
neighboring ranches, because the control pastures under study were isolated
(>10 km. separation), with primarily agricultural crops (citrus and dates)
in the areas between. Two principal
predators present were Philonthus
discoideus Gravenhorst
and Philonthus
longicornis
Stephens. Their abundance in control
pastures was significantly greater on all collection dates than where O. gazella populations were firmly established. Another staphylinid, Platystethus spiculus
Erichson, was present under both situations, but this species is probably not
an obligate predator, preferring to feed on manure (Legner & Moore,
1977). Populations of ants
and predacious mites were also present, but not monitored. Other predatory species in the Histeridae, Carabidae and Cincindelidae only infrequently were found at very low
densities. Reasons For Continued Adult Fly
Abundance.--The continued abundance of adult horn flies
on cattle suggests that a predicted 38-56% reduction of hornflies in O. gazella pastures was insufficient to noticeably reduce the
adult fly density congregating on single animals. However, emigration of excess flies from these pastures could
have been reduced so that an area wide reduction of hornflies might have occurred. Nevertheless, the reduction from a
supersaturated to a saturated environment did not obviously give a noticeable
level of control on cattle, as judged with binocular observations at 10-11 AM
4-5 PM. A similar situation might
prevail in Australia where imported scarab activity seems sufficient to cause
significant reductions in bush fly, M.
vetustissima, breeding but
which paradoxically is not accompanied by drops in the annoyance thresholds. Explanation For Staphylinid Reduction.--The lower numbers of Philonthus spp. in irrigated pastures where O. gazella were highly active may be found in the dynamics of
scarab beetles with horn flies and their natural enemies in the dung
habitat. Natural enemy habitats are
undoubtedly altered or destroyed by the dung shredding process. The shredding activity of O. gazella reduces habitat configuration and moisture content
to a level that may be unsuited for staphylinid oviposition and larval
development. In some respects this is
similar to the effects of cultivation on the natural breeding habitat of Hippelates eye gnats, which
causes a marked reduction in the effectiveness of natural enemies (Legner
& Olton, 1969).
Other evidence that scarabs
in America may disturb Philonthus species was given by Roth et al. (1983) who associated declines in these predators'
abundance with rising scarab population densities. Although the
introduction of additional scarab beetle species may afford a positive means
for lowering exophilic fly densities, it is important to consider whether
introduced scarabs might, through habitat disruption, preclude the
introduction of effective predatory species.
Because there are no practical nonbiological control methods to reduce
fly numbers in exophilic habitats, and the addition of more scarabs may
actually exacerbate the problem, the most logical direction for research is
to intensify worldwide searches for more effective natural enemies,
especially predators and pathogens. REFERENCES: [Additional references may be found at: MELVYL
Library ] Anderson, J. R. & E.
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