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CONTROL OF THE BUSH FLY, MUSCA
SORBENS WIEDEMANN & OTHER
DIPTERA IN KWAJALEIN ATOLL, MARSHALL
ISLANDS (1) E. F. Legner (2), B.
B. Sugerman (3), Hyo-sok
Yu (4), and H. Lum (5) (Contacts) Please CLICK on
underlined figures, tables & footnotes to view details] SUMMARY An investigation into the biological-integrated
control of pestiferous flies in Kwajalein Atoll, Marshall Islands revealed 6
principal species in the families Calliphoridae, Sarcophagidae and Muscidae,
of which were successfully reduced to generally non-annoyance levels in 18
months. The 6th species, Musca sorbens Wiedemann, was satisfactorily
reduced on some islands through habitat reduction, the introduction of
natural enemies and baiting after 21 months. The history, biology and control
of M. sorbens is reviewed and future efforts outlined for the importation of
natural enemies. possibilities for resistance to insecticides are indicated
and precautions recommended for the effective long term use of poisoned
baits. The potential invasion of M. sorbens in North America is discussed. (7) INTRODUCTION Pestiferous
flies in the Marshall Islands provide a classic example of the adaptation of
invading noxious insects to a salubrious tropical climatic area. With nearly perfect
temperature-humidity conditions for their development, an abundance of
carbohydrate and protein-rich wastes in the form of garbage and excreta
provided by man and his animals, and a general absence of effective natural
enemies, several species were able to reach maximum numbers. There are
principally 4 types of pestiferous flies in Kwajalein atoll of the Marshall
Islands, with the African-Australian bush fly, Musca sorbens
Wiedemann, being by far the most pestiferous species (FIG. I). The common house fly, Musca demestica L., of
lesser importance, frequents houses and is attracted to food in recreation
areas. The remaining 2 types are the Calliphoridae [Chrysomya megacephala
(Fab.), and Phaenicia cuprina (Wiedemann)) , and the Sarcophagidae [ Parasarcophaga
misera (Walker), and Phytosarcophaga gressitti Hall and Bohart).
These latter species are abundant around garbage disposal sites and wherever
rotting meat and I decaying fish are available. The Calliphoridae are copper
to bluish-green, sometimes blue, on the thorax making them easily
distinguishable from the large striped gray Sarcophagidae. Both of these
flies differ from the common house fly and the bush fly in being more
sluggish and noisy and by their general avoidance of humans. Because
residents do not a distinguish different kinds of flies, these latter 2 types
are often blamed as nuisances when in fact they may be considered to fulfill
a useful role in the biodegradation of garbage and rotting a meat. PROCEDURE Studies
were begun in April 1971 at the request of the United States Army and the
United States Trust Territory of the Pacific, to investigate fly abatement
possibilities in Kwajalein Atoll, Marshall Islands. The atoll, the largest in
the world, is about 120 Km long by an average 23 Km wide and contains 93
islets grouped roughly between 166-167° E. longitude and 8-9° N. latitude in
an irregular ring around a central lagoon (Bryan 1972 and FIG. 2). Kwajalein, the
largest island, lies at the extreme south of the atoll (FIG. 2 , 3 & 5). An initial
assessment of the problem using standard sampling methods to locate fly breeding sources and identify
associated natural enemy fauna, resulted in the expedient implementation of
breeding source reduction to reduce the house fly, Musca domestica L., and
both the Calliphoridae and Sarcophagidae to inconspicuous levels, which
largely involved slight modifications of refuse disposal sites to disfavor
fly breeding. By November 1972, these simple measures resulted in an
estimated 1/3rd reduction of total flies concentrating around beaches and
other recreational areas (see Table 1 for proportion of fly species trapped). Because the house
fly especially enters dwellings, its
reduction was desirable for the general health of the community, and fly
annoyances indoors diminished.
Thorough surveys of breeding sites and natural enemy complexes
revealed that Musca sorbens reduction would not be.quickly
forthcoming, however. A schedule of importation for natural enemies was begun
in 1971 and other integrated control approaches were investigated: eg.
baiting and breeding habitat manipulation.
The following account of this, the principal problem fly species in
the atoll, reviews bush fly history and biology , at the same time pointing
out the magnitude of the biological, technological and political difficulties
involved in its permanent reduction. RESULTS AND DISCUSSION Bush Fly. Musca sorbens Wiedemann 1830 Origin and
Habits.
-- This species is known as the bazaar fly in North Africa, a house fly in India,
and the bush fly in Australia (Yu 1971). It was first described from Sierra
Leone in West Africa in 1830 (Wiedemann 1830) where it is a notorious
nuisance to man and animals. The flies are attracted to wounds, sores, and
skin lesions, searching for any possible food sources such as blood and other
exudations. Although not a biting species, its habits of being able to
transmit eye diseases, entheric infections, pathogenic bacteria and helminth
eggs make it one of our most important and dangerous public health insects
(Bell 1969; Buston and Hopkins 1927; Gaud and Faure 1951; Gaud et al. 1954;
Greenberg 1971; Hafez and Attia 1958; Lamborn 1937; McGuire and Durant 1957;
Patton 1920, '22, '23, '26, '32, '33; Thomson and Lamborn 1934). The bush fly
has spread through a major portion of the Old World, Africa and parts of Asia
(Awati 1920; Gaud and Faure 1951; Hafez and Attia 1958; Hughes 1968; Jack
1931; Lewis 1954; Meng and Winfield 1944; Norris 1966; Patton 1920, '23, '26;
Roy and Siddoils 1940; Van Emden 1965). In Oceania its distribution is in AustraIia (Norris 1966;
Paterson and Norris 1970); New Guinea (Paterson and Norris 1970); Samoa
(Huxton and Hopkins 1927); Guam (Harris and Down 1946, Hohart and Gressitt
1951); and the Marshall Islands (Hohart and Gressitt 1951 )0 In Hawaii it was
first reported by Joyce in 1950. Since then Hardy (1952) listed it in the
Catalog of Hawaiian Diptera, and Wilton (1963) reported its predilection for
dog excrement. Annoyance by the bush
fly increased in the 1960's when it was incriminated as a potential vector of
Beta-haemolytic streptococci in an eipdemic of acute glomerulonephritis (Bell
1969). Taxonomy. -- Paterson and Norris
(1970) identified 3 biological forms
in the M. sorbens complex. These are distinguished by morphometric
differences and reproductive isolation in the laboratory , and consist of the
African male broad-frons and narrow-frons forms, and the Australian
narrow-frons form. Examination
of morphometric measurements of 220 randomly selected adult males collected on Kwajalein Island during
the period April 1971 -November 1972 showed an average frons ratio (6) of 0.0590 (range 0.0341 -0.1052; SD = 0.0182), which was
close to that of both the African and Australian narrow-frons forms. However,
the frequency distribution showed 2 distinctive peaks representring the
narrow- and broad-frons in the ratio of about 4: I (FIG 4). Also, a count of the parafrontal hairs of
20 random females (Paterson and Norris 1970) gave similar results. Thus, 2
forms may be sympatric in Kwajalein, which has been shown possible from other
areas (Paterson and Norris 1970). Breeding
Habits.
-- The principal breeding sites of the bush fly vary according to different
geographical localities. However,
human excrement is regarded by many authorities as a preferred site when this
is available (Bohar,t and Gressitt 1954; Gaud et al. 1954; Hafez and Attia
1958; Harris and Down 1946; Norris 1966; Patton 1933; Peffly 1953a, b; Roy
and Siddons 1940; Sabrosky 1952). The
bush fly also breeds in fresh droppings of a number of larger animals, such
as cattle (Bohart and Gressitt 1951; Hafez and Attia 1958; Hughes and Walker
1970; Jack 1931; Meng and Winfield 1944; Pat ton 1922, '36; Peffly 1953b);
horses (Bohart and Gressitt 1951; Lewis 1954; Pat ton 1922, '36); dogs (Meng
and Winfield 1944; Norris 1966; Pat
ton 1922; Wilton 1963); and pigs (Bohart and Gressitt 1951; Meng and Winfield
1938; Smirov 1940; Hafez and Attia 1958). Sheep feces (Norris 1966), and goat
and cat feces (Yu 1971) are also suitable.
Poultry excrement will not support this fly's development (Yu
1971). Other breeding sources of much
lesser importance but which are capable of producing small numbers of bush
fly are carcasses of animals, decaying vegetable, matter and garbage (Bevan
1926, Norris 1966, Pat ton 1922, Yu 1971).
Recent studies by Yu (1971) in Hawaii show that dog feces are the most
important breeding medium on Oahu. He
concluded that dung of cows, horses, pigs and goats are of minor importance
in rural areas where dog feces are not readily available. Development. -- The average
developmental period in Hawaii from eggs to adults is 10 days with an average
of 76 flies emerging per field sample of dog feces. On Kwajalein
developmental time probably averages 8 days with over 100 flies emerging from
one sample of dog feces. High precipitation may prevent the fly from full
activity and breeding (Yu 1971). In the
Kwajalein atoll there is little reason to doubt that a substantial portion of
the main density of Musca sorbens emanates from dog and human
feces. However, extensive inspection
of pig droppings in the bush of 10 widely separated islets revealed high
numbers of larvae ( over 100 per dropping), making this dung, as in Guam
(Bohart and Gressitt 1951 ), a primary breeding source in the atoll. Pigs
that are corralled on soil or concrete slabs concentrate and trample their
droppings making them less suitable breeding sites. In such situations flies
were only able to complete their development along the periphery of
corrals. Coconut husks placed under
pigs in corrals results in the production of greater numbers of flies by
reducing the effectiveness of trampling. These conclusions were reached after
comparing quantitative samples taken over a period of 2 years from the
respective breeding habitats. Garbage was
not found to breed M. sorbens, although admittedly a very low
percentage of the adult population could originate there judging from reports
elsewhere. Nevertheless, this medium is certainly not responsible for
producing more than 2% of the adult densities observed in the atoll. Control
Efforts and Evaluation Worldwide. -- Successful partial control of bush fly has been
achieved only in Hawaii through a combination of the elimination of breeding
sites, principally dog droppings, and the activities of parasitic and
predatory insects introduced earlier for biological control of other fly
species, e.g., Musca domestica (Legner 1978 ). The density of-bush fly
varies in different climatic zones in
Hawaii, but the importance of this fly is minimal compared to Kwajalein. At
times hymenopterous parasites have been found to parasitize over 95% of flies
sampled in the Waikiki area (H. Yu, unpublished data). Other parts
of Oceania.. were either not suitable for the maximum effectiveness of known
parasitic species ( e.g. Australia) or the principal breeding habitats were
not attractive to the natural enemies. Therefore, in Australia a concerted
effort is being made to secure scavenger and predatory insects from Africa
that would be effective in the principal unmanageable fly producing source,
cattle and sheep dung (Bomemissza 1970; G. F. Bomemissza, personal
communication). For example, insectary studies on Onthophagus gazella
F. (Coleoptera: Scarabaeidae) showed 80-100% reduction of M. sorbens
emergence from cattle dung (Bomemissza 1970). Surviving fly larvae gave rise
to small, stunted adults of reduced reproductive capability. Kwajalein
Atoll.
-- Continuing integrated fly control begun in 1971 has now reached a level of
partial success. Initial surveys for natural enemies of M. sorbens in
April-May 1971 revealed the presence of 4 scavenger and predatory insects,
the histerid Carcinops troglodytes Erichson, the nitidulid Carpophilus
pilosellus Motschulsky, the tenebrionid Alphitobius diaperinus
(penzer), and the dermapteran Labidura riparia (pallas). Following our
recommendations in May 1971, dogs were significantly reduced and all privies
were reconstructed or improved on one island, Ebeye Dogs were reduced or tethered on Kwajalein Island and refuse
fish, etc., disposed of thoroughly on l1leginni (an island that experienced
outbreaks of Sarcophagidae in 1970 (FIG. 2),
and other islands with American residents.
Continuing importations of natural enemies were made throughout the
atoll with establishment of some species verified in November 1972 (Table 2). The average
density of M. sorbens on Ebeye was, subsequently, considerably reduced by
November 1972 from an estimated 8.5 flies attracted to the face per minute!,
to less than 0.5 flies per minute, a reduction readily recognized by the
inhabitants (7). The single most important cause appeared
to be the partial elimination of breeding sources, with natural enemies
playing a secondary role The M.
sorbens density at Roi-Namur (estimated 2.5 flies attracted per minute), in
the norther apex of the atoll, and Ennylabegan (6.0 flies per minute), in the
southwest, did not substantially improve following the natural enemy
introductions and recommendations for reducing breeding habitats. However, this is thought to be due to an
increase in the number of human inhabitants, dogs and pigs on Ennylabegan and
on Ennubira to the southeast of Roi-Namur (Fig. 2). Flies
apparently invade islands adjacent to their breeding origin at a very slow
rate as evidenced by a lack of recovery on Kwajalein of dyed flies that were
released on Ebeye and Ennylabegan in November, 1973. However, Australian
studies support the fact that M. sorbens possesses rapid,
long-distance migrating behavior, which is aided by windstorms (Hughes 1970,
Norris 1966). Even though mark and recapture experiments were negative, there
is probably continuous slow invasion of M. sorbens over a proposed
route from Ennubira to Roi-Namur; and from Ennylabegan to Enubuj and
Kwajalein (FIG. 2 & FIG. 3). Flies are
absent on islands without native or American inhabitants even though adjacent
islands may have high densities, which lends some support to the minimal
invasion hypothesis. Fly persistence around
human habitation is probably not to seek breeding sites but rather to
attain foodstuffs, especially carbohydrates, meat scraps and sauces which are
perennially available at recreation sites especially. Therefore, control efforts stress
corralling pigs, reconstructing privies and reducing dog populations. All
residents on all islands are urged to refrain from indiscriminate disposal of
wastes from soft drink and beer cans. Importations of natural enemies is
being stressed for those areas where pigs and dogs roam wild. New
Approaches to Bush Fly Reduction. -- New approaches to the solution of the bush fly
problem in Kwajalein atoll involve the use of effective poisoned baits for adult
flies, technological changes in garbage disposal and the importation of
natural enemies from the presumed original endemic area of M: sorbens in
central and eastern Africa. Poison Baits Sugar bait
mixtures that have been used for M domestica in years previous to 1972
were wholly ineffective for killing adult M. sorbens due to their
almost complete lack of attractiveness.
Baits tested during November 1972 through April 1973 included a
variety of decomposing foodstuffs, of which rotting eggs and rotting fish
sauces were very highly attractive. In one experiment using a 6-day old
mixture of one part fresh whole eggs to one part water (Legner 1970b, 1971), it is estimated that about
50,000 bush flies were attracted to the mixture and killed by a 0.5 ppm
Dichlorvos, (8) additive. The poisoned mixture was poured in
quantities of 100 mI each on damp sand at 20 sites in the shade and spaced
every 10m along a public beach on Kwajalein (FIG
6 & FIG 7). Baits placed above the height of 1m or against walls in
open pavilions were only weakly attractive. After 48 hours, flies were
reduced to inconspicuous levels all over Kwajalein Island (Table 1 gives the
approximate % of each species trapped).
This condition endured for 3 consecutive days after which immigrating
flies managed to build up to annoying levels starting on the 4th day when the
baits ceased to be attractive. The former density of flies had not yet been
reached, one week after the baiting experiment; and this population was
subsequently reduced in the same manner by applying additional fresh poisoned
baits. Flies breeding in dog feces were the principal source of the
population recovery on Kwajalein, as shown by marking and recovery
experiments in November 1973.. Baiting was
continued through November 1973 and extended to other islands in the atoll
with a result of sustained reduction of bush fly to below general annoyance
levels (less than 0.01 attracted per minute)1O on Kwajalein, Roi-Namur,
Illeginni and Meck Islands. A new
attractant that augmented the rotting egg mixture consisted of beach sand
soaked for one week in the decomposing body fluids of buried sharks. This new
attractant was far superior to rotting eggs both in rate and time of
attraction, the latter sometimes exceeding 5 days. The baiting
method of control can be used effectively if applied initially twice a week.
A schedule of biweekly applications or longer may follow as determined by
personnel in charge of the control
program. Resistance Resistance
in the tropical environment could be expected to develop very quickly if
adult flies that are exposed to the baits are able to reach breeding sites.
This resistance might ultimately be expected to spread through the whole M.
sorbens population in Kwajalein atoll, similar to that observed for other
species (Georghiou 1966). Obviously, if breeding sites and poisoned baits are
located together within the normal flight range of flies, resistance rate is
increased. However, there is probably a lower resistance possibility to
electric shock devices, and not much chance for resistance against a natural
bait such as rotting eggs or fish.
Nevertheless, with full awareness of the plasticity of M sorbens and
the general resilience of nature, it cannot be ignored that the possibility for resistance to any
chemical control implemented by humans does exist. Technological and Cultural
Improvements To further ensure against breeding of M.
sorbens around refuse disposal sites and that resistance does not develop
in populations of the common house fly and blowfly which are able to
successfully breed in these sites,
the development of suitable concrete barriers to larval flies around such
sites was proposed. Fly larvae of M. domestica and Phaenicia
cuprina especially, were repeatedly observed entering the soil from cans
and dumpsters containing refuse. Such refuse sites placed on adequately
rimmed concrete platforms would probably prevent larvae from finding suitable
pupation sites in surrounding soil. The use of petroleum oils in a long
perimeter around refuse deposition sites, saturating the soil within the
depicted circle to a depth of at least 5 cm, might offer partial control.
However, some fly larvae are capable of penetrating this barrier so that its
effectiveness is not as great as concrete. Natural
Enemies Biological
control organisms are usually effective only in one or a few breeding
habitats, and their activities are limited under certain types of climatic
stress. For example, when house flies breed in garbage they are less prone,
to attack by certain natural enemies than when breeding occurs in accumulated
animal excrement (Legner and Poorbaugh 1972). The aim of biological
control isto locate and establish natural enemies that will perform
effectively in: all problem breeding habitats. This goal is rarely achieved,
so that effective control takes on an integrated aspect where cultural and
chemical means are implemented. There are
no known natural enemies that will specifically attack the bush fly nor is there
any information on what kinds of parasites and predators range in the
droppings of humans, dogs and pigs in eastern Africa, where bush fly may have
originated. The best that can be done in biological control at this time is
to import natural enemies of other fly species for their possible adaptation
to M. sorbens. (Legner 1970a; Legner and Greathead 1969 ; Legner and Olton 1968, 1970, 1971 ; Legner et al. 1967). This is the
approach that has been made in Hawaii and is partially successful. As new species of natural enemies become
available, they will be introduced for biological control. There is some
expectation that a concerted effort will be made to seek out specific natural
enemies in eastern Africa as support for the work becomes available. A list
of species that should be considered
for introduction in Kwajalein atoll is shown in Table 3; however, this list is expected to grow as information
about new natural enemies from Africa develops. The
biological control phase of fly reduction must be viewed as a long range
approach with ever increasing fly reduction as the new beneficial species
become established. An investment of research time in the initial evaluation
of a natural enemy species during the importation phase would not be
practical from the standpoint that such time would, (I) detract from the
amount of effort placed on introductions; (2) most species will probably not
become established judging from a history of biological control efforts during
the past 50 years; and (3) fly densities are expected to drop gradually with
the relatively small numbers of natural enemies being released at any given time. Once an obvious drop in the
adult M. sorbens density has occurred, a thorough evaluation of the
natural enemy species responsible may be carried out. POTENTIAL INVASION OF MUSCA SORBENS IN NORTH AME RICA There has
been some speculation about the possibility of Musca sorbens invading
portions of North America, where climatic similarities with Africa and
Australia exist. Undoubtedly the more tropical portions of the continent
could sustain this species, with the Florida peninsula and Mexico appearing
the most vulnerable. There is greater uncertainty about its establishment in
the Southwestern United States where
winter cold may be an effective barrier. In Australia M .sorbens over
winters with difficulty in the colder southern quarter of the continent
(Norris 1966) and probably reinvades that region annually from the more
tropical north (Hughes 1966). Similarly, in the Middle East Israel seems to
be out of the general range of M. sorbens (Legner and Olton 1968), although it is a prominent
species further south along the Nile River (Hafez and Attia 1958). There
being no sustained breeding sites available in the intervening desert,
invasion during warm seasons is precluded. Much the same conditions exist in
the American Southwest where annual invasions from Mexico, a potential
breeding source, may be minimal across the arid Sonoran Desert. The threat of
invasion in the southeastern United States from potential breeding sites in
Florida is greater, however. CONCLUSIONS In modern
pest control the importance of being conscious of what effects a control
action may have on the total environment as well as its permanency are
recognized. Resistance to pesticides, especially in the tropics, is a greater
threat in our procedure than pesticide pollution because we employ compounds
that degrade and are nonpoisonous to us. In fly control on Kwajalein,
resistance to Dichlorvos and related compounds would leave no alternative
stop-gap measure to use in case of an epidemic of human disease in which
flies were vectors. Resistance could
also produce hardier strains and theoretically even more aggressive flies (G.
P. Georghiou, personal communication; Legner
1970a). Therefore, it is essential that the baiting
program to combat flies is minimized and that stress be placed on the implementation
of technological, cultural and biological controls that could lead to a
permanent reduction of fly breeding in the atoll. As of
January 2002, the baiting procedure in Kwajalein has not continued with the
sophistication initially determined necessary. The result is that flies are not adequately reduced. Personnel changes that resulted in
critical information not being passed on are primarily responsible. Of paramount importance is the proper
brewing of the baits and the latter’s placement wind calm areas of the
islands. Also, the baiting program
should logically be extended, at least experimentally, to Musca
vetustissima Walker in Australia, where towns simulate islands in the
vast bush land. Temporary relief from
bush fly annoyance might be expected to occur during summer when these flies
descend into the higher latitudes. ACKNOWLEDGMENTS The
assistance and cooperation of personnel in the following organizations who
made this scientific study possible is gratefully acknowledged: Global Associates, Department of the Army,
University of California, United States Trust Territory of the Pacific, and
the people of the Marshall Islands. REFERENCES CITED
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s. Thesis, Univ. of Hawaii. 72 p. --------- ---------- (1) This study was supported in part by
grants- in-aid and assistance of Global Associates, Oakland, California. (2) Professor of Biological
Control, University of California, Riverside 92521. (3) Entomologist, Trust Territory of The
Pacific (4) Research Assistant, University of
California, Riverside 92502. (5) Foreman, Facilities, Global Associates,
Kwajalein, APO San Francisco 96555 (6) Ratio of the least width of the frons and
the greatest width of the head. (7)
Estimates of Musca sorbens adult density were made by the senior
author between 10-1500 hours by counting the number of alightments per minute
on the face, neck and ear areas of the body while standing upright in the
shade on the rainless days. At least 20 such estimates were gathered on
anyone island per day and the results averaged for text presentation.
sources, with natural enemies playing a secondary role. (8) 2,2-dichlorovinyl dimethly phosphate |
