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The arthropod community in pastures and its biocontrol potential for the horn fly, Haematobia irritans (L.) in north-central Florida

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The arthropod community in pastures and its biocontrol potential for the horn fly, Haematobia irritans (L.) in north-central Florida
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Hu, Guangye, 1954-
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Arthropods ( jstor )
Beetles ( jstor )
Cattle ( jstor )
Eggs ( jstor )
Feces ( jstor )
Insects ( jstor )
Larvae ( jstor )
Pastures ( jstor )
Predators ( jstor )
Species ( jstor )
Greater Orlando ( local )

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THE ARTHROPOD COMMUNITY IN PASTURES AND ITS BIOCONTROL
POTENTIAL FOR THE HORN FLY, Haematobia irritans (L.)
IN NORTH-CENTRAL FLORIDA



















By

GUANGYE HU


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA


1995














LD
1760
1~
42O

















ACKNOWLEDGMENTS

I wish to express my deepest gratitude to my major

professor, Dr. J. Howard Frank, for his invaluable guidance,

encouragement, advice and financial support. Thanks are

also extended to Drs. J. F. Butler, L. P. Lounibos, J. A.

Hogsette, R. S. Sand for serving on the supervisory

committee and contributing to the completion of the

dissertation. I wish to extend special thanks to Dr. Butler

for his help with photographing insects, allowing me to use

the equipment at his laboratory, and providing me access to

his horn fly colony for my experiments, and Dr. Lounibos for

loaning me a video camera for my research.

I am also grateful to Mr. P. Dixon and J. Stokes for

granting me use of the beef pastures of IFAS, University of

Florida, for my field research. I wish to acknowledge Dr.

D. Williams for the help with application of Amdro for fire

ant population control, Dr. T. Fincher for donating an

emergence box designed to extract insects from cowpats, and

Dr. J. Castner for photographing emergence and testing

equipment used for this study.

I would like to thank the following specialists for

helping me with the insect identification: K. Ahlmark, F.

ii














Bennett, P. Choate, G. A. Dahlem, D. Deonier, R. J. Gagnd,

W. L. Grogan, V. Gupta, D. Habeck, M. E. Hennessey, T. J.

Henry, P. W. Kovarik, G. W. Kranz, J. McNamara, F. Mead, A.

S. Menke, K. Nguyen, M. E. Schauff, P. Skelley, A. Smetana,

R. J. Snider, G. Steck, G. B. Edwards, J. Watts, M. Thomas,

D. Williams, W. Wirth.

I am appreciative of members of the insect biocontrol

lab, including J. Cicero, R. Coler, R. Hemenway, P. Parkman

and S. Wineriter for helping me and creating a comfortable

and friendly working atmosphere; D. Simon, H. Brown and J.

Okine in Dr. Butler's lab for helping me to collect immature

horn flies and offering me fly chow essential for the

experiments on predation.

Special thanks go to my family. During the four years'

study my wife, Yanfen Chen, endured and sacrificed much

(especially when she was pregnant with our son) to ensure

time for my study. I appreciate my daughter, Wenli, who

exercised a great deal of tolerance and understanding of my

absence.


iii















TABLE OF CONTENTS


ACKNOWLEDGMENTS ..........

LIST OF TABLES ..........


ii


vii


LIST OF FIGURES .


. . ix


ABSTRACT

CHAPTERS


1. INTRODUCTION


. 1


2. LITERATURE REVIEW ....

History and Importance of
History .....
Economic Importance
Biology of the Horn Fly
Life History ....


the Horn Fly


Mating and Oviposition .. ........
Host Orientation and Location . .
Diapause and Dispersal .. ........
Control of the Horn Fly ... .........
Chemical Control ... ..........
Genetic, Physical and Immunological
Control ..... ............
Biological Control .... ........
Dung Arthropod Community .......
Dung Arthropod Community Composition
Succession of the Community . .
Effect of the Whole Insect Fauna on
Horn Fly Production .........
Predators . . . . . . .


Staphylinidae .... .......
Hydrophilidae .... .......
Histeridae ......
Red imported fire ant .......
Mites ..... .............
Flies ..... ............


Competitors ............
Coprophagous beetles .....
Coprophagous flies ......


.4

4
4
.4
.7
7
* 10

13
14
14


Q
O


g


I
t


6








Parasitoids ............
Ecology and Biology of Staphylinidae

3. The ARTHROPOD COMMUNITY IN NORTH CENTRAL
FLORIDA PASTURES ............


Materials and Methods
Results and Discussion
Coleoptera ....
Staphylinidae
Scarabaeidae
Carabidae
Hydrophilidae
Histeridae .
Diptera .........
Hymenoptera ...
Other Insects and An


* . 34
* . 35


* . 38


imals .. ......


4. EFFECT OF ARTHROPOD PREDATORS ON HORN FLY
SURVIVORSHIP IN PASTURES AND UNDER LABORATORY
CONDITIONS ........ ................ 73

Materials and Methods ... .......... 74
Test of the Arthropod Community on
Horn Fly Survivorship .... 74
Laboratory studies on horn fly
survivorship .......... 74
Field studies on horn fly
mortality .. ....... 75
Determination of Predation Rates of
Predators ... ........... 77
Collection and colonization of
predators .... ..... .77
Predation rate test under
laboratory conditions . 77
Predation rate test under
simulated field conditions 78
Results and Discussion .... ......... 82
Laboratory Horn Fly Survival . . 82
Fauna Caused Horn Fly Mortalities 82
1992 July trial .. ........ 82
1992 August trial ....... 83
Predation Rates of Predators on Horn
Fly Immatures ... ......... 84
Philonthus spp ... ........ .84
Under laboratory conditions 86
Under simulated field
conditions .. ..... .92
Oxytelinae .... ....... 96
Paederinae ... .......... 100
Xantholininae .. ........ 100
Tachyporinae ... ........ 105








Aleocharinae .. ......... 105
Hydrophilidae .. ....... 107
Histeridae .... ...... 107
Carabidae ..... ........ 108
Tenebrionidae ..... ...... 109
Anthicidae ..... ......... 109

5. EFFECT OF THE RED IMPORTED FIRE ANT (RIFA) ON THE
HORN FLY AND OTHER ARTHROPODS IN PASTURES . 111

Materials and Methods .. ......... 112
Results ........................ 115
RIFA Population Control .. 115
RIFA Effect on the Horn Fly . . 117
RIFA Effect on Epigeal Arthropods 122
1992 trial .. .......... 122
1993 trial .. ......... ..122
RIFA Effect on Dung-Inhabiting
Arthropods ... ......... 124
1992 trial ... ......... 124
1993 trial .. .......... 127
Discussion .... ............. 134

6. CONCLUSION AND GENERAL DISCUSSION ...... ..137

REFERENCE LIST ......... ................... 146

BIOGRAPHICAL SKETCH ....... ................. 170














LIST OF TABLES


Table


Page


3-1. Summary of invertebrates collected in pastures in
north-central Florida from 1991 to 1993 ...... ..42

3-2. Staphylinidae collected in pastures in north central
Florida ....... ..................... 45

3-3. Scarabaeidae collected in pastures in north central
Florida. ........ ...................... 53

3-4. Coleoptera collected in pastures in north central
Florida, with exclusion of Staphylinidae and
Scarabaeidae ...... ................... 58

3-5. Diptera collected in pastures of north central
Florida ......... ...................... 62

3-6. Hymenoptera collected in pastures in north central
Florida ......... .................. 66

3-7. Miscellaneous invertebrates collected in pastures in
north central Florida ..... ............... ..68

4-1. Larval predation rates of five Philonthus species
during the whole developmental period on horn fly
eggs and larvae under laboratory conditions . . 91

4-2. Predation by N. pusillus larvae and adults on eggs
and larvae of horn flies under laboratory
conditions ........ ................... 102


5-1. Numbers of horn flies emerged from artificial pats
in the Amdro-treated and control areas, October
1992 and 1993 . . . . . . . . .

5-2. Numbers of arthropod specimens collected by pitfall
traps from the Amdro-treated and control areas in
October 1992 .... ........... .........


vii


119


120


120










5-3. Numbers of staphylinid specimens collected by
pitfall traps from the Amdro-treated and control
areas in October 1993 ..... .............. 121

5-4. Numbers of arthropod specimens collected by pitfall
traps from the Amdro-treated and control areas in
October 1993 ....... ................... 125

5-5. Numbers of staphylinid specimens collected by
pitfall traps from the Amdro-treated and control
areas in October 1993 ..... .............. 126

5-6. Numbers of specimens of arthropod taxa collected
per cowpat sample from the Amdro-treated and
control areas, October 1992 ... ........... 129

5-7. Numbers of specimens of selected arthropod taxa
collected per cowpat sample from the Amdro-treated
area and the control area, October 1993 ..... ..131

5-8. Numbers of staphylinid specimens extracted from
Amdro treatment and control area in October 1993 133


viii


Table


Page














LIST OF FIGS


Figure Page

3-1. Emergence boxes used for extracting arthropods from
intact cowpats of 24 hrs old ... ........... 41

3-2. Seasonal distribution of five families of Coleoptera
most commonly collected by trapping .. ........ .43

3-3. Seasonal distribution of species diversity of
Staphylinidae and Scarabaeidae in pastures of north
central Florida ....... .................. 44

3-4. Seasonal distribution of staphylinid species in
pasture A, collected by trapping ... ......... 47

3-5. Seasonal distribution of scarabaeid species in pasture
A, collected by trapping ..... .............. ..54

4-1. Cone traps used for covering cowpats to collect horn
and other flies emerged from the pats. . ... 79

4-2. Cages used for testing predation rates of
predators on horn fly immatures .... ......... .80

4-3. Horn flies emerged from covered and uncovered cowpats
in a north central Florida pasture in 1992 . . 85

4-4. P. longicornis adult preying on horn fly eggs under
laboratory conditions ..... ............... .88

4-5. Adult predation rates of five species Philonthus on
horn fly eggs and larvae per day under laboratory
conditions ........ ..................... .89

4-6. Daily predation rates of 5 species Philonthus larvae
on horn fly eggs and larvae under laboratory
conditions ....... .................... 90

4-7. Predation rates by four species of Philonthus on horn
flies under simulated field conditions .. ....... ..94








Figure


Page


4-8. Predation of P. lonaicornis on horn flies at
different prey densities under simulated field
conditions ........ .................... 95

4-9. Daily predation rates of N. pusillus larvae on horn
fly eggs and larvae under laboratory conditions 103

4-10. Aleochara notula preying on horn fly eggs under
laboratory conditions ..... .............. 104

5-1. Fire ants preying on horn fly larvae
in the laboratory ...... ................ 118














Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment
of the Requirements
for the Degree of Doctor of Philosophy

THE ARTHROPOD COMMUNITY IN PASTURES AND ITS BIOCONTROL
POTENTIAL FOR THE HORN FLY, Haematobia irritans (L.)
IN NORTH-CENTRAL FLORIDA


By

Guangye Hu

May 1995

Chairperson: J. Howard Frank
Major Department: Entomology and Nematology

A complete arthropod community was investigated by

pitfall traps and emergence boxes from 1991 to 1993. Over

60,000 invertebrates were collected in pastures. Arthropods

found were 226 species belonging to 73 families in 14

orders. Coleoptera were the most diverse as judged by

trapping and extracting (109 species), Diptera the second

(35 species) and Hymenoptera the third (24 species). Most of

the beneficial insects were in these three orders.

Field mortality of horn flies caused by the arthropod

community was tested by seeding horn fly eggs underneath

artificial cowpats, which were either exposed or isolated

from the arthropod community by using cone traps. The

community-caused mortalities of horn flies in the field were

75.9% and 66.7% in July and August 1992, respectively. The

xi








coverall average was 71.3%.

Predation rates were tested under laboratory (Petri

dishes) and simulated field conditions (test cages

containing artificially formed pats) with adults of five

field-collected Philonthus species against horn fly

immatures. P. longicornis had the highest predation rate.

Aleochara notula was very abundant during the survey and its

adult is an effective predator of horn fly immatures.

Fifteen other staphylinid, one hydrophilid, one histerid,

and two carabid species were also found to prey on horn fly

eggs and larvae. Larvae of eight staphylinid species and

one tenebrionid species were also found to prey on horn fly

immatures.

Red imported fire ants (RIFA) were observed to infest

cowpats heavily all year round and prey on horn fly larvae

and pupae in the pastures. RIFA caused 94.3% and 62.9%

mortality of horn flies, respectively, for October 1992 and

1993, compared with a RIFA-controlled area in the field,

though it reduced populations of horn fly natural enemies.


xii














CHAPTER 1
INTRODUCTION


The horn fly, Haematobia irritans (L.), is an Old World

species which invaded the United States before 1886. Its

populations grew and spread, and reached Florida by 1891.

Its range is from Canada to Brazil (Hargett and Goulding

1962b; Moya-Borja 1990) and it can be a serious problem for

the cattle industry in the New World wherever it occurs.

Adult male and female flies are hematophagous, and bite

frequently (Harris et al. 1974). Cattle are their primary

hosts and fly numbers on a single animal can reach thousands

(Bruce 1964; Butler 1975, 1990; Fay 1986; Kinzer 1970). The

blood loss and annoyance can cause substantial reduction of

milk production and live weight gain in domestic cattle

(Bruce and Decker 1947; Harvey and Brethour 1979).

The fly larvae are coprophagous and develop only in

fresh cattle dung (Hogsette and Koehler 1986; Macqueen and

Beirne 1975; Skidmore 1991). Studies have shown that other

insects present in cattle dung are important mortality

factors for the immature stages of the horn fly (Blume et

al. 1970; Fay 1986; Fay and Doube 1983; Fay et al. 1990;

Harris 1981; Harris and Blume 1986; Kunz et al. 1972;

Legner 1986; Macqueen and Beirne 1975; Sanders and Dobson










1969; Thomas and Morgan 1972b). The horn fly viability is

affected by predation, parasitoidism, and competition for

the same food source (Harris and Blume 1986). Important

horn fly predators are reported to be Staphylinidae,

Hydrophilidae, Histeridae (Harris and Blume 1986; Thomas and

Morgan 1972b), and the red imported fire ant (Howard and

Oliver 1978; Schmidt 1984; Summerlin et al. 1984b). The most

important competitors are dung beetles (Scarabaeidae)

(Fincher 1986; Anderson and Loomis 1978; Blume et al. 1973).

The most abundant parasitoids are Spalangia spp. (Harris and

Blume 1986).

Although the parasitoids of horn flies have been

investigated in Florida (Butler et al. 1981; Escher 1977),

the arthropod community in cattle dung has not been examined

quantitatively, and little attention has ever been paid to

its species composition. Such information, however, is very

important for evaluation of the biotic mortality of horn fly

immature stages, and for determination of the need for

introduction of biocontrol agents for horn fly control.

The primary goal of the present study was to provide

extensive information on the species composition and

abundance of horn fly natural enemies and their effect on

the horn fly immatures in north central Florida. In detail,

the objectives of this study were 1) to determine the

composition and abundance of the arthropod community in

cattle dung; (2) to monitor seasonal distribution of these








3
arthropods; 3) to determine the effect of the whole

arthropod community on horn fly production from cattle dung;

4) to determine the species of predatory beetles in cattle

dung and evaluate their predation ability on horn flies; 5)

to determine the effect of the red imported fire ant (RIFA)

on horn fly production in pastures; and 6) to study the

biology of the predatory Staphylinidae found in cattle dung.














CHAPTER 2
LITERATURE REVIEW


History and Importance of the Horn Fly


History

The horn fly, Haematobia irritans (L.), is a

widespread, economically important pest of cattle (Morgan

and Thomas 1974, 1977). It hitchhiked into the United States

from southern France before 1886, but before then was known

only from Europe (Riley 1889). This fly is believed to have

entered the United States on cattle through the port of

Philadelphia and was first collected in Camden, New Jersey,

in August 1887 (Bruce 1964; Morgan and Thomas 1974). It

spread rapidly in North America (Hargett and Goulding 1962b;

Stone et al. 1965; Butler 1990), reaching Florida by 1891

(Osborne 1896). It reached South America later, and is

still moving south (Moya-Borja 1990). This fly has occupied

almost the whole of Brazil, Paraguay, Uruguay and Bolivia,

and over half of Argentina (Foil and Hogsette 1994).



Economic Importance

Horn flies are conspicuous because they can occur on

the host in large numbers. It has been estimated that

10,000 flies may occur on a single animal (Bruce 1964).

4








5
Bulls often have larger numbers than cows. Young calves

seem to be less bothered than either cows or bulls. Horn

fly adults in Florida reach 10,000-20,000 flies per animal

with bimodal peaks in abundance in May-June and August-

September (Butler 1990).

The horn fly is one of the most damaging pests of

cattle, especially in southern areas where fly populations

may reach several thousands per animal during the long fly

season (Kinzer 1970). Damage caused by the horn fly is due

to irritation of the host, loss of blood, reduced vitality,

and refusal to graze when the numbers are high (Bruce 1964).

The repeated biting of hundreds to thousands of flies

daily irritates cattle. The energy extended in efforts to

dislodge the flies, including tail switching, head slinging,

walking and aggregation (Harvey and Launchbaugh 1982),

causes unmeasured losses. Both sexes of the adults suck

blood. Daily feeding frequencies are reported as two or a

few times (Bruce 1942, 1964) to 20 times (Koehler and Butler

1980), or even more frequently (Harris et al. 1974). Meal

size taken per fly varies from 0.045 to 0.19 mg. Feeding

occurs throughout the day. In the laboratory, Harris et al.

(1974) observed that females fed up to 38 times a day, with

an average of 163 minutes feeding per day; males fed 24

times per day, with an average of 96 minutes feeding per

day. A population of 10,000-20,000 flies per animal would

cause a loss of 2 liters of blood per month (Butler 1975).










Dollar losses to the cattle industry due to horn flies

are increasing. The annual losses were estimated as three

million in 1942 (Bruce 1942), 150 million in 1956 (Knipling

and McDuffie 1956), 179 million in 1976 (Steelman 1976), 730

million in 1981 (Drummond et al. 1981), and 870 million in

1991 (Kunz et al. 1991), respectively. In Florida, Koehler

and Butler estimated losses were $36 million in 1976 and $50

million in 1984, respectively; Butler later estimated a

loss of about $61 million in 1986 (Hogsette and Koehler

1986).

The losses include the reduction of weight gain, or

weight loss in cattle (Cheng 1958; Cutkomp and Harvey 1958;

Roberts and Pund 1974; Campbell 1976; Harvey and Brethour

1979; Laake 1946), as well as reduction in milk production

in dairy cattle (Bruce and Decker 1947; Granett and Hansens

1956, 1957). Weight gains of 30 to 70 pounds were observed

after animals were treated with DDT (Laake 1946). Grannett

and Hansens (1957) showed a decrease in milk production of

one-fourth in unprotected dairy herds. Bruce (1947) showed

a high inverse correlation between changes in milk

production and fly abundance.

The damage threshold of fly numbers per cow was

reported to be from 50 (Butler 1975) to 200 (Hogsette et al.

1991, Schreiber et al. 1987b), or even higher (Haufe 1982).

The heterogeneity of these data is caused by the

experimental design, the region where the tests were








7

conducted, the breed and sex of animals, and the size of the

horn fly population (Steelman 1976).

Horn flies serve as vectors for a filarial nematode,

Stephanofilaria stilesi, and a bacterial disease, mastitis,

between cattle. The nematode reduces the value of hides

for leather and causes blemishes that are a problem when

registered animals are used for exhibition purposes

(Steelman 1976). Mastitis may render dairy cattle useless.

Horn flies are also potential vectors of other diseases

(Butler 1975).



Biology of the Horn flV

Life History

The horn fly belongs to the Animal Kingdom Arthropoda

-Atelocerata (Uniramia) Hexapoda (Insecta) Pterygota -

Diptera Aristocera Muscoidea Muscidae Haematobia

irritans (Linnaeus) (Borror et al. 1989). It is a

holometabolous insect, with four stages: egg, larva, pupa

and adult. After the host has been found, the adult horn

fly leaves the vicinity of the host only to oviposit on

fresh manure. Eggs hatch within 24 hours and the larvae

pass through three instars, pupating in the third instar

skin in 3 to 5 days. It takes nine days from oviposition to

adult emerge at 300C (Melvin and Beck 1931). The field

developmental time depends on the temperature (Hargett and

Goulding 1962a; Lysyk 1992a&b; Wilkerson 1974).










Eq. Most of the eggs laid are reddish and difficult

to detect in manure (Hogsette and Koehler 1986), but a small

percentage of tan, yellow, and white eggs is also laid

(Morgan and Schmidt 1966). The egg is 1.2 mm long and 0.32

mm wide (Miller et al. 1984) and is usually deposited on the

undersurface of the edge of fresh dung pats. The embryo

hatches from the egg less than 24 hours after deposition in

the field during the summer, but embryonation time depends

upon temperature (Melvin and Beck 1931; Melvin 1934; Depner

1961). Temperature and moisture conditions affect the egg

development significantly (Bruce 1964; Lancaster and Meisch

1986; Wilkerson 1974).

Larva. The horn fly larva is cream-colored, with the

anterior pointed and the posterior truncate. It has a mouth

hook at the front, a pair of 4-6 branched anterior spiracles

and a pair of heavily pigmented knob-like posterior

spiracles. The larva passes through three instars in the

manure within four days (Wilkerson 1974). The newly hatched

larvae immediately seek a crack, crevice, or perforation in

the manure into which they crawl to obtain food and shelter.

When the manure pat crust becomes firm and dry, horn fly

larvae move to a more suitable part of the manure (Bruce

1942, 1964). About 92.5 hours after hatch, larvae migrate

for pupation to the underside of the manure or into the soil

depending on the relative moisture content of the

alternatives (Bruce 1964; Escher 1977).








9
Under laboratory conditions horn fly larvae can develop

in cattle, bison, sheep, and horse manure (Greer and Butler

1973), but in the field larval development takes place only

in cattle manure (Depner 1961, Bruce 1964, Foil et al.

1990). Experiments show that cattle fed a large amount of

grain produce manure which is unusually acid, reducing horn

fly larval survival under laboratory conditions (Schmidt

1983).

Pupa. The pupa is coarctate and the puparium is dark

brown. It is approximately 3.3 mm long and 1.4 mm wide. This

stage lasts about 5.5 days under natural field conditions.

It can differ by three to four days depending whether the

manure pat is in sunlight or shade (Kunz et al. 1970).

Emergence of the flies from manure occurs from mid to late

afternoon in the summer, but at any time of day in the

spring and fall (Lancaster and Meisch 1986). Males need one

day less to develop at this stage than do females (McLintock

and Depner 1954; Hoelscher and Comb 1971). Sex ratio was

reported as 1:1 or 1:1.35 (Glaser 1924; Mohr 1943).

Adult. The adult horn fly is about 4 mm long (Foil et

al. 1990), or about one-third to one-half the size of the

common house fly, Musca domestica (L.) (Okine 1991).

Longevity of the horn fly is estimated from 28 days

(McLintock and Depner 1954) to 6-8 weeks (Bruce 1964). Male

survival rates were reported to be about 95% of the female

survival rate (Krasfur and Ernst 1983). Both sexes have










piercing and sucking mouthparts and are blood feeders.



Matinq and Oviposition

Mating

Adults began to mate one day after eclosion under

natural conditions (Harris et al. 1968), but two or three

days after eclosion under laboratory conditions (Bruce 1964,

Lancaster and Meisch 1986). Females are monogamous, but

males can inseminate an average of 4.6 females in the field

(Harris et al. 1968).

Mating behavior has been described by Bolton (1980) and

Zorka and Bay (1980). Mating behavior of the male involves

orientation, tapping the dorsum of the female's abdomen with

prothoracic tarsi, mounting and using its legs to grasp the

female, positioning genitalia towards the female's, grabbing

the female's ovipositor with claspers, and copulation.

Copulation lasts from 0.9 to 5 minutes (Zorka and Bay 1980),

3.5 to 11.5 minutes (Bolton 1980), and 0.5 to 5 minutes

(Bruce 1964). Mating is chemically mediated and males are

attracted by cuticular lipids of the female (Mackley et al.

1981).
Ovipiosition

Females begin laying eggs one day after mating (Bruce

1942, 1964; Harris et al. 1968; Schmidt et al. 1972) and

oviposit at any time during the day and night (Sanders and

Dobson 1969; Kunz et al. 1970). Horn fly eggs are deposited










only on fresh cattle droppings (Butler 1975; Kunz et al.

1970; Sanders and Dobson 1969; Skidmore 1991). Droppings

older than 10 minutes are unattractive unless the crust is

broken (Bruce 1964). The flies move from the shoulders of

cattle to the area near the tail,,as animals deposit manure

(Foil and Hogsette 1994). Possibly the gravid female

receives a signal of impending defecation with the lifting

of the tail of the host, and is ready to oviposit on the

fresh manure as soon as it hits the ground, or even before

the feces hit the ground (Skidmore 1991). Mohr (1943)

observed that all adult horn flies left the dung pats two

minutes after the dung was deposited. Each female lays an

average of 24 eggs per batch and 15 batches during its

lifetime (Bruce 1964). The eggs are laid singly or in

groups of 4 to 6 (Hogsette and Koehler 1986), and a total of

200 to 400 is laid during the life time of the female

(Wilkerson 1974).



Host Orientation and Location

Adult flies remain on the body of cattle day and night,

unless the females leave for oviposition (Hammer 1941;

Hargett and Goulding 1962b; Kunz et al. 1970; Morgan

1964), though they make short rapid flight from one part of

the host to another, or between adjacent hosts (Bruce 1964).

The name 'horn fly' originates from the tendency of these

flies to cluster at the base of horns of the host, but










clustering of flies on horns is no longer as common as in

previous times. (Foil and Hogsette 1994). The flies

congregate mainly on the shoulders and sides of the animal

(Hogsette and Koehler 1986), because these places are least

disturbed by the swishing of the tail, tossing of the head

and kicking of the legs (Harvey and Launchbaugh 1982). Horn

flies alter their distribution (from back and shoulder area

to belly area) on cattle treated with pyrethroid-impregnated

ear tags in comparison with untreated cattle (Byford et al.

1987).

In a series of experiments, Hargett and Goulding

(1962b) found that many horn flies, dislodged during the

night, failed to find the host until daylight. They

concluded that upon emerging, horn flies depend more on

vision than on olfactory or heat stimuli to find a host. In

contrast, Kinzer et al. (1978), using an artificial cow as a

field attractant, found that temperature and CO2 were prime

factors in horn fly orientation. Dalton et al. (1978) found

that radiated heat rather than internal temperature was

involved and that cow odor was the most powerful influence

on responding flies, especially at close range.

Color and breed are reported to affect the horn fly's

host preference. Horn flies prefer dark-colored animals

(Bruce 1964). Brahman cattle were less attractive than

European breeds (Tugwell et al. 1969). Calves and yearlings

with long, thick hair are not as desirable as are cattle








13
with short, thin hair (Hammer 1941). Bulls attract and hold

more horn flies than cows, because of the effect of male sex

hormone, testosterone (Dobson et al. 1970). Though the horn

fly's principal host is cattle, it sometimes attacks other

animals such as goats, horses, mules, deer, dogs and,

rarely, man, especially when the bovine host is absent

(Bruce 1964).



Diapause and Dispersal

Hargett and Goulding (1962b) reported that the horn fly

overwinters in the southern United States as an adult, but

in the northern United States and Canada it overwinters as a

diapausing third instar larva or as a pupa. There has been

no observed diapause in horn flies in Florida (Wilkerson

1974). Horn fly diapause is mediated in the fall by

decrease of temperature and photoperiod (Thomas et al. 1987)

and is terminated in the spring by increasing temperatures

of the substrates (Lysyk 1992b; Thomas et al. 1987).

Some authors stated that horn fly adults are sedentary

on the host, but leave for oviposition on manure pats (Bruce

1942, 1964; McLintock and Depner 1954). On the contrary,

Chamberlain (1981, 1982) and Hoelscher et al. (1968) found

that horn flies moved several hundred meters from a host.

Moreover, Kinzer and Reeves (1974) and Tugwell et al. (1966)

showed the potential for long distance dispersal of natural

horn fly populations. Horn flies were found to be able to








14
disperse 5 km or even further (Sheppard 1994). The movement

occurs nocturnally (Hoelscher et al. 1968) and dispersing

populations are predominantly females (Marley et al. 1991).



Control of the Horn Fly



Chemical Control

Control of horn flies with chemical insecticides has

been the primary method for the last century and it is

widely practiced today. Methods of insecticide application

include sprays, dust bags, ear tags, tail tags, leg bands,

and back rubbers for killing the adults, and feed-through

for killing fly larvae (Beadles et al. 1975; Butler and

Koehler 1979; Drummond et al. 1988; Hogsette and Koehler

1986; Schreiber et al. 1987a). Beginning with the

widespread use of DDT and related chlorinated hydrocarbon

insecticides, and later with organophosphorus compounds,

horn flies were controlled effectively (Morgan and Thomas

1974, 1977). The advent of insecticidal ear tags (Byford et

al. 1985; Harvey and Brethour 1970, 1979; Quisenberry and

Strohbehn 1984; Schmidt and Kunz 1980) permitted highly

effective season-long horn fly control in most regions.

In the USA, insecticide resistance in horn flies

occurred sporadically before the 1980s when horn flies were

easy to control with the available organochlorine and

organophosphate insecticide. Two to three years after the










pesticide-impregnated ear tags were in use, widespread

insecticide resistance occurred throughout the country

(Sheppard 1990). Subsequent development of horn fly

resistance to other insecticides, especially pyrethroid

compounds (Quisenberry and Strohbehn 1984; Sheppard 1984;

Sheppard and Joyce 1994; Kunz and Schmidt 1985), has

decreased field efficiency of chemical insecticides used for

horn fly control. In addition, insecticides used for feed-

through for control of dung-inhabiting flies leave a residue

in cattle dung, which adversely affects abundance of

biocontrol agents (e.g. Scarabaeidae and Staphylinidae) of

the horn fly in pastures (Fincher 1992; Madsen et al. 1990;

Wall and Strong 1987).



Genetic, Physical and Immunological Control

Sterile male technique has been used for horn fly

control (Eschle et al. 1977; Kunz and Eschle 1971) and was

successful. The difficulties for keeping this technique

continue to work are isolation of the target area (Kunz et

al. 1974), raising large numbers of competitive sterile

flies, and difficulty in combining several control measures

(Graham and Hourrigan 1977).

A practical horn fly walk-through trap designed by

Bruce (1938) has been tested to control the horn fly. This

trap controlled an average of 54-73% and 50% horn flies on

cattle in Missouri (Hall and Doisy 1989) and Texas (Bruce










1938), respectively. An updated version is being tested as

the University of Maryland (Hogsette, pers. comm.).

Less attractive hosts, such as Brahman (zebu) or

Brahman-cross cattle (Tugwell et al. 1969; Steelman et al.

1994) have been reported to decrease horn fly populations.

However, weight gain with these cattle are usually less than

those of English breeds (Steelman et al. 1994).

Studies show that exposure of animals to hematophagous

arthropod ectoparasites, such as ticks, mosquitoes and horn

flies, evokes an immune response to the parasite (Kerlin and

Allingham 1992). Developing a vaccine would enhance the

host immune response which would be deleterious to

arthropods' feeding on the host. Vaccines are possibly

developed to inoculate cattle against horn fly bites in the

future.



Biological Control

Though research emphasis on horn fly control in North

America has been centered on using chemical insecticides,

increasing attention has been given in the past 2-3 decades

to an examination of the field relationships and

interactions between immature stages of the horn fly and the

other organisms that share its habitat.

The horn fly is a serious pest in North America, but

not in its origin, Europe, because the insect fauna of

droppings is reportedly richer in Europe than in North










America. Staphylinidae, for example, have been considered

the most important predators in cattle dung. Fourty-three

species were found to be associated with cattle dung in this

country (Fincher 1990), while 133 species were reported from

cattle dung in a study in Finland (Koskela 1972). The horn

fly is also an immigrant pest in Australia and has plagued

the cattle industry there, but it rarely reaches pest status

in Africa (Zumpt 1973), because there is a higher level of

fauna-induced mortality in Africa than in Australia

(Bornemissza 1960; Fay 1986). Natural enemies inhabiting

cattle dung in Europe, Africa and Asia have been introduced

for horn fly control in North America (Fincher 1990; Merritt

and Anderson 1977). Introduction of additional biocontrol

agents for biocontrol of the horn fly is needed (Fincher and

Morgan 1990; Fincher 1990; Fincher and Summerlin 1994),

because arthropods currently inhabiting cattle dung have not

suppressed the horn fly populations to an acceptable level

in North America (Fincher 1990; Hunter et al. 1991).



Dung Arthropod Community



Dung Arthropod Community Composition

Basic information on the arthropod fauna of dung is

essential for consideration of introduction of biocontrol

agents for horn fly control. Bovine manure supports a large

population of insects, mainly including many dipterous,










coleopterous and hymenopterous species. Information from

North America shows that arthropod species that affect horn

flies vary geographically, but overall groups are generally

the same (Harris and Blume 1986). On the other hand,

information from South Africa shows that the dung faunal

species composition and abundance differ between habitats,

although a degree of overlap exists (Fay 1986). Each

habitat has its own distinct community of dung organisms

that are effective control agents of horn flies. There is

only a small core of common species that occur in all the

habitats and these play the major role in maintaining the

fly population at a certain level (Fay 1986). Most of the

beneficial insects belong to the orders Coleoptera, Diptera

and Hymenoptera (Fincher 1990; Harris and Blume 1986) with

many other arthropods of less importance (Sanders and Dobson

1966). Mohr (1943) was one of the first workers in the

United States to consider the entire insect complex of

cattle droppings. Since then, several comprehensive surveys

have been conducted throughout the country (Macqueen and

Beirne 1975; Merritt and Anderson 1977; Sanders and Dobson

1966; Poorbaugh et al. 1968; Blume 1970; Valiela 1969b;

Wingo et al. 1974). Studies on specific groups of the

arthropods in pastures in America north of Mexico were

summarized by Blume (1985), who included a checklist,

distribution maps, and an annotated bibliography. Over 400

species of arthropods have been collected in or on cattle










dung in the United States.

Mohr (1943) reported 67 species of insects inhabiting

dung, representing 28 families of 5 orders in Illinois.

Poorbaugh et al. (1968) reported 151 species of insects

attracted to and reared from cowpats in California,

representing 76 families in 4 orders. Sanders and Dobson

(1966) reported 38 insect species including 17 families of 3

orders in Indiana. Blume (1970) reported 103 insect

species representing 45 families of 5 orders in Texas.

Macqueen and Beirne (1975) reported 67 arthropod species

representing 21 families of 4 orders in British Columbia,

Canada. Valiela (1969b) reported 109 species of insects and

mites in 43 families of 7 orders in New York. Cervenka and

Moon (1991) reported 108 arthropod species in 19 families of

4 orders in Minnesota. Wingo et al. (1974) reported 157

arthropod species in 33 families of 5 orders in Missouri.



Succession of the Community

Members of the newly forming dung arthropod community

start to arrive immediately, with the horn fly prominent

among the first colonists. Many other species arrive during

the first few minutes such as Sarcophagidae (Mohr 1943). And

the major period of colonization lasts until a distinct

crust of the dung has formed (Skidmore 1991). Hunter et al.

(1986a) observed that arrival time of Staphylinidae ranged

from 30 minutes to 24 hours. The insects arriving within








20

the first 24 hours are very important for reducing the horn

fly populations (Fay et al. 1990; Blume et al. 1970).



Effect of the Whole Insect Fauna on Horn Fly Production

The presence of other arthropods in cattle dung has

been shown to reduce populations of horn flies (Blume et

al. 1970; Fay 1986; Fay and Doube 1983; Harris and Blume

1986; Kunz et al. 1970, 1972; Macqueen and Beirne 1975; Roth

1989; Sanders and Dobson 1969; Thomas and Morgan 1972b).

Blume et al. (1970) showed a significant negative

correlation between the mean numbers of horn flies and the

mean numbers of insects of other species produced per dung

pat. Roth (1989) reported that dung fauna caused an average

of 87.9% mortality in immature horn flies in naturally

infected dung pats between April to mid-September in Texas.

In a similar study in Missouri, Thomas and Morgan (1972b)

reported 97.7% mortality of horn flies in exposed dung pats.

When dung pats were covered within five minutes to eliminate

competition from other insects, an average of 66.5 flies

emerged from each pat (Kunz et al. 1970); but when pats were

not covered and competition from other insects was allowed,

an average of 6.6 flies emerged per pat (Kunz et al. 1972).

In Florida, Wilkerson (1974) and Greer and Butler (1973)

found an average of 8 to 19 flies emerging from each manure

pat in the summer months.

In Australia, Fay et al. (1990) reported that fauna-










induced mortality of horn flies was from 79% to 84% and

fauna-reduced headcapsule width of the emerged horn flies

was an average of 2% to 7%. Small flies produced by

indirect effect of the dung fauna are less fecund and have

lower survival potential (Fay 1986; Roth 1989).



Predators

Non-arthropod animals, especially birds, have been

considered predators of flies (Hammer 1941), but research

attention has been focused on arthropods inhabiting cattle

dung. Studies in Missouri (Thomas and Morgan 1972b), Texas

(Blume et al. 1970; Roth 1983, 1989; Roth et al. 1983), and

Canada (Macqueen and Beirne 1975) indicated that predators

are the primary insect biotic mortality factors affecting

the horn fly.

Studies have shown that the most important predators

are coleopterans of the families Staphylinidae,

Hydrophilidae, and Histeridae (Bornemissza 1968; Fay 1986;

Fay and Doube 1983; Fincher 1990; Legner 1986; Macqueen and

Beirne 1975; Roth 1989; Roth et al. 1983; Sanders and Dobson

1969; Thomas and Morgan 1972b), especially Staphylinidae

(Hammer 1941; Blume et al. 1970; Macqueen and Beirne 1975;

Thomas and Morgan 1972b).

Staphylinidae. Staphylinidae are the most important

predators because of species diversity and high population,

and predation rate (Fincher 1990; Harris and Blume 1986).










Fincher (1990) reported 43 species associated with cattle

dung in the continental USA. The important subfamilies

include Aleocharinae, Oxytelinae, Paederinae, Staphylininae

and Tachyporinae (Hunter et al. 1991). Staphylinid beetles

prey on the dung fauna as adults and as larvae (Drea 1966;

Fincher and Summerlin 1994). Predation by Staphylinidae on

horn fly eggs and early instar larvae has been reported for

Philonthus (Harris and Oliver 1979; Hunter et al. 1989;

Macqueen and Beirne 1975; Roth 1982; Thomas and Morgan

1972b), and Aleochara (Klimaszewski and Blume 1986; Harris

and Blume 1986).

Members of the genus Philonthus are important predators

of dung-inhabiting flies as adults and larvae (Hammer 1941;

Laurence 1954; Sanders and Dobson 1966; Valiela 1969a;

Macqueen and Beirne 1975; Wingo et al. 1974; Wharton 1979).

Fifteen species of Philonthus were reported to inhabit

cattle dung in this country (Fincher 1990). P. cruentatus

(Gmelin) was shown to be one of the most effective predators

on immature stages of the horn fly (Thomas and Morgan

1972b). P. flavolimbatus Erichson was shown to reduce horn

fly production by 69% and 86 when 2 and 4 beetles were

tested for each trial (Roth 1982). Horn flies were reduced

by 91% and 99% when 5 and 10 beetles of this species were

tested for each trial (Harris and Oliver 1979). P.

flavolimbatus and P. cruentatus were found to prey

primarily on the egg stage, while P. rectangulus Sharp is a










predator of larvae (Roth 1982). Three exotic species, P.

longicornis (cosmopolitan), P. flavocinctus (from southeast

Asia) and P. minutus (from Australia), were tested to reduce

horn fly emergence by 61.1%, 68.1% and 99.2%, respectively,

under laboratory conditions (Fincher and Summerlin 1994).

Eight species of Aleochara are known to occur in cattle

dung in the USA (Fincher 1990) and the adults prey on the

eggs and larvae of cyclorrhaphous Diptera. But in the

immature stages they are solitary ectoparasitoids of fly

pupae within the host puparium (Klimaszewski and Blume

1986).

Hydrophilidae. Sixteen species of hydrophilid beetles

have been reported from cattle dung in the continental USA

(Fincher 1990). Members of the genus Cercyon are abundant,

but the adults and larvae are dung-feeders (Sanders and

Dobson 1966; Hafez 1939; Merritt 1976; Thomas and Morgan

1972b). Sphaeridium scarabaeoides (L.), which is very

common in dung, has been considered as an important horn fly

predator in larval (Bourne and Hays 1968; Hammer 1941;

Macqueen and Beirne 1975; Mohr 1943; Poorbaugh et al. 1968;

Sanders and Dobson 1966; Thomas and Morgan 1972b) and adult

stages (Hammer 1941). The larva showed the highest predation

rate on horn fly larvae at 800F, and no predation below 40'F

under laboratory conditions (Bourne and Hays 1968).

Histeridae Twenty-two species of histerid beetles are

associated with cattle dung in the continental USA (Fincher










1990) and members of this family have been reported to be

predators of developing dipterous larvae in cowpats

(Bornemissza 1968; Thomas and Morgan 1972b; Summerlin et al.

1981, 1984c, 1990; Wang et al. 1990). Laboratory tests

(Summerlin et al. 1982, 1984c, 1990) have indicated that

Hister coenosus Erichson is the most effective histerid

species in reducing horn fly populations. It reduces horn

fly populations by 99.1%, when the adult beetles are exposed

to all immature stages of the horn fly. Next in order of

effectiveness are Hister incertus Marseul (98.7%), H.

abbreviatus F. (96%), Atholus rothkirchi Bickhardt (55.8%),

Saprinus pennsylvanicus (Paykull) (40.9%) and Xerosaprinus

orbiculatus (Marseul) (40.8%). Predation rates increase as

the density of predators increases (Summerlin et al. 1982,

1990). In Texas, Summerlin et al. (1989, 1991) also found

Phelister panamensis LeConte, P. haemorrhous Marseul, and

Pachylister caffer Erichson to be predators of horn fly

immatures.

Red imDorted fire ant. Other species of ants, such as

Pogonomyrmex californicus (Buckley) (Wharton 1979), are

reported as very important predators of horn flies, but the

most emphasizes the red imported fire ant (RIFA), Solenopsis

invicta Buren. This ant arrived near Mobile, Alabama,

approximately 50 years ago from South America and has spread

across the southern United States (Porter and Savignano

1990). RIFA has been labelled a serious agricultural pest










that damages crops and attacks livestock, poultry and

wildlife (Hays and Hays 1959; Lyle and Fortune 1948). RIFA,

however, has been reported to prey on dung-inhabiting

immature Diptera (Laurence 1954; Bruce 1964). It invades

fresh dung pats which are less than 10 minutes old

(Summerlin et al. 1984a) and causes significant reduction of

horn flies in field studies (Howard 1975; Howard and Oliver

1978; Summerlin et al. 1984b; Schmidt 1984). Howard and

Oliver (1978) and Schmidt (1984) observed that fire ants

carried larvae and pupae of the horn fly from the pats.

Summerlin et al. (1977) showed that RIFA reduced emergence

of adult horn flies about 20 fold from bovine feces in a

laboratory study in Alabama; Schmidt (1984) showed a seven-

fold increase of the horn fly in a pasture in Texas where

RIFA was controlled. Howard and Oliver (1978) found that 2-

5 times more horn fly pupae were recovered from pasture in

Louisiana where RIFA was controlled by Mirex bait than in

pasture where it was not controlled. Lemke and Kissam

(1988) reported that the number of horn flies emerging from

manure piles where RIFA was controlled using Pro-Drone was

55% greater than the number that merged from piles in a

RIFA-infested field in South Carolina.

On the other hand, some evidence shows that RIFA not

only affects pest fly populations but also has negative

effects on some other beneficial insects inhabiting the

dung, such as the scarab beetle Onthophagus gazelle F.










(Summerlin et al. 1984b) in Texas and Staphylinidae in

Louisiana (Howard and Oliver 1978)i

Mites. Several species of miftes encountered in dung

are predators of flies. These mites are in the following

families: Parasitidae, Uropodidae, Eviphididae, Laelapidae,

Pachylaelapidae, and Macrochelidae (Krantz 1983).

Macrochelidae, in particular, have been considered to have

potential for biocontrol of dung-inhabiting flies (Anderson

1983; Doube et al. 1986; Krantz 1983; Axtell 1963) and have

been shown to be efficient predators of house flies (Axtell

1963; Krantz 1983), bush flies, face flies and horn flies

(Halliday and Holm 1987; Anderson 1983). These mites

commonly appear on the surface of droppings as soon as

beetles arrive, because many are mainly carried phoretically

by dung-inhabiting beetles including Scarabaeidae (Mohr

1943; Poorbaugh et al. 1968; Krantz 1983; Stewart and Davis

1967), Trogidae and Histeridae (Stewart and Davis 1967).

Halliday and Holm (1987) tested nine species of

macrochelid mites as predators of the bush fly and horn fly.

The mites preyed on fly eggs and larvae, but preferred

larvae over eggs. Macrocheles pere rrinus (Krantz), M. qlaber

(MUller) and M. peniculatus Berlese were the most efficient.

In Australia, Macrocheles pererinus has been imported

from Africa and established (Roth et al. 1988b). This

species was shown to attack horn fly eggs in all stages of

development, and to kill the larvae 24 hours after hatching.








27

Each mite has the capacity to kill 8-10 flies when there is

a high density of eggs provided. It caused 66% and 54%

suppression of horn fly populations, respectively, in two

field tests (Doube et al. 1986). In another experiment,

however, M. peregrinus caused an average of 33% suppression

of horn flies in field cowpats, and had a stronger

preference for eggs of other dipterous species that have

softer chorions (Roth et al. 1988b).

Flies. Although some 200 species of Diptera have been

reported on or in cattle dung, only the flies that develop

in the dungpats can contribute to the reduction of the pest

flies by competition and predation. Members of the families

Muscidae, Empididae and Sarcophagidae have been considered

to be possible predators (Harris and Blume 1986). The flies

reported to be predacious are Drapetis spp. (Empididae)

(Laurence 1952), Hydrotaea (Muscidae) (Merritt 1976; Hammer

1941), Myospila meditabunda (F.) (Muscidae) (Poorbough et

al. 1968), Gymnodia (=Brontaea) spp. and Orthelia spp.

(Muscidae) (Ferrar 1975) and Ravinia lherminieri (Robineau-

Desvoidy) (Sarcophagidae) (Pickens 1981). Hammer (1941)

reported that Hydrotaea larvae are typically facultative

predators. Data on suppression of horn fly populations by

these species are unavailable. Myospila meditabunda

(Muscidae) was assumed to inflict high mortality on horn fly

and other coprophagous larvae (e.g. sarcophagids,

scatophagids, sepsids, and sphaerocerids) in California










(Poorbaugh et al. 1968).



Competitors

A horn fly larva needs approximately 2 mg dung to

complete its development (Macqueen and Beirne 1975). A

single adult bovine drops an aver ge of 12 dung pats every

day (Waterhouse 1974), yielding a total weight of some 30

kilograms of feces. This amount of manure can support

15,000 flies (horn and stable flies are the main pests of

cattle in the USA). Moreover, if they are not disposed of,

the pats produced by each animal will blanket between 5 and

10 percent of an acre in a year. In addition, at the

periphery of each dung pat, there develops a zone of tall,

rank herbage that cattle seldom eat and avoid for a year or

more unless they are ravenous. The effective area of

pasture is thereby reduced by each bovine by about one-fifth

of an acre per year (Waterhouse 1974). There are

approximately 10.5 million cattle in the United States

(Campbell 1993), producing more than 120 million pats a day.

Without competitors, accumulated manure pats produce

trillions of pest flies and cause loss of large areas of

pasture.

Coprophagous beetles. Dung beetles (Scarabaeidae) have

long been regarded as useful agents in the control of flies

that develop in dung of domestic animals (Fullaway 1921;

Lindquist 1936). Since the horn fly lays its eggs in fresh








29

dung, survival from eggs to adult flies should be reduced if

dung deposits are rapidly buried or eaten by dung beetles

(Fincher 1986). There are approximately 5,000 species of

Scarabaeidae worldwide (Skidmore T991). Certain species in

the subfamilies Aphodiinae, Scarabaeinae, and Geotrupinae

are usually abundant (Hanski 1991; Woodruff 1973; Skidmore

1991). Most scarab species belong to the subfamily

Scarabaeinae, which contains 4,000 species (Bornemissza

1976).

Coprophagous beetles cause mortality of horn flies by

competing for the same food source (cattle dung), and by

moving and burying dung to reduce and interrupt larval

habitats of the horn fly (Andersor and Loomis 1978; Blume et

al. 1973; Bornemissza 1970, 1976; Doube and Moola 1988;

Ferrar 1975; Fincher 1981, 1986, 1990; Waterhouse 1974).

These beetles also reduce dung accumulation, improve

pastures by increased fertility and improved soil structure

(Fincher 1981, 1990; Fincher et 1. 1983). Rapid removal

of feces would return areas of pasture to grazing (which

normally would be lost because of contamination), recycle

tons of nitrogen normally lost into the atmosphere, and

reduce pest fly populations on li vestock. Dung tunnelers

also provide runways so predatory staphylinids can get at

the flies (Valiela 1974). The larvae of dung beetles also

digest bacterial albumens which may account for their

subsistence in old dung heaps (Merritt 1976). The benefit








30
yielded by scarabs results in pol ential savings of some two

billion dollars in the USA (Finchnr 1981).

The effect of dung beetles on suppression of horn flies

has been best shown in Africa, where upward of 2,000 species

of coprid beetles are known to us the dung of many species

of herbivorous vertebrates (Waterl ouse 1974). Therefore,

the horn fly is not a problem to t he African cattle

industry. In Britain, Skidmore (2.991) reported that

Aphodius contaminatus descend on cowpat and disrupt other

arthropod colonists. Sometimes A. contaminatus is so

abundant that there may be more beetles than dung and the

effects of such visits are to scatter the dung over a wide

area and render it useless for other community members.

Using coprophagous scarabs to reduce populations of

dung-inhabiting arthropods has rec-eived much attention

(Anderson and Loomis 1978; Blume et al. 1973; Bornemissza

1970, 1976; Fincher 1981, 1986). There have been active

programs for the importation and establishment of exotic

coprophagous scarabs in Australia (Bornemissza 1976) and the

United States (Blume et al. 1973).

Although more than 100 specie s of dung beetles have

been reported from cattle dung on pastures in the U. S.,

millions of cattle droppings remain on the surface of

pastures for several months (Fincher 1990). This is

because native dung beetles and other coprophagous organisms

cannot effectively consume and re ove them (Fincher 1986,








31
1990). Hence, further introduction of dung beetles is needed

in this country.

Fullaway (1921) introduced dung beetles in Hawaii as a

biological control agent for the horn fly. Twenty-three

species of dung beetles were introduced in Hawaii from 1906-

1963 (Fincher 1986). During the 1970s, several additional

species were released in Hawaii by courtesy of the CSIRQ

Dung Beetle Project from Australia. Experiments in Hawaii

in 1966 showed 95% fewer horn flies emerged from cowpats

attacked by Onthophagus gazella than from pats from which

these beetles were excluded (Bornemissza 1970). So far, 15

species of dung beetles have been released in the

continental USA (Fincher 1990). The following species have

been established: 0. taurus Schreber, which was released in

Texas and established in southern States (Fincher and

Woodruff 1975; Fincher 1990) and California (Anderson and

Loomis 1978); 0. gazella, which was released in Texas

(Blume and Aga 1978), California and Georgia in 1975,

Arkansas in 1976 and Mississippi in 1979 (Fincher 1981) and

established in southern States (Fincher et al. 1983; Hunter

and Fincher 1985); 0. depressus Harold which was established

in Florida and Georgia by unknown means (Fincher 1990);

Euoniticellus intermedius Reiche, which was released and

established in Texas (Blume 1984); Onitis alexis Klug,

which was released in Texas and established in California

(Anderson and Loomis 1978).








32

Bornemissza (1960) advocated the use of certain species

of dung beetles in Australia to effect disposal of surface

dung, improve pastures by incorporation of dung into soils,

and reduce larval habitats for the horn fly. The CSIRO

Division of Entomology began a program to import foreign

dung-burying scarab beetles into ustralia in 1964

(Bornemissza 1976). The purpose of the project was to reduce

the populations of the horn fly in north Australia

(Waterhouse 1974; Bornemissza 197 ) by increasing the

natural dispersal of cattle dung. By 1991, CSIRO had

imported and released 55 species cf dung beetles in

Australia (Doube and Macqueen 199 ). Colonized scarab

species cause high mortality of hcrn flies and other

dipterans when their populations are high (Roth et al.

1988a, 1991).

Many species of Hydrophilida7 and Staphylinidae, which

are not usually referred to as dung beetles, also use some

components of the decomposing material or the microorganisms

in dung pats (Hanski 1991). Whether they suppress the horn

fly production has not been specifically reported, but

Merritt (1976) and Valiela (1969b, 1974) reported that

Sphaeridium burrowed in and out of the dung shortly after

pats were dropped, providing aeration to the pat and

permitting staphylinids and parasitic Hymenoptera to use

their tunnels to locate their tar ets.

On the other hand, the compatibility of scarabs with










the predators for horn fly biocontrol is questionable.

Large numbers of scarabs burrowing through the dung could

cause significant dung removal an pasture improvement

(Fincher 1981; Legner 1986; Waterl ouse 1974) but might not

achieve significant fly reductions (Legner 1978, 1986;

Macqueen 1975). High numbers of dung beetles could disrupt

the oviposition behavior of female staphylinids and reduce

the food source of immatures, causing a decrease of the

numbers of predators (Legner and Narkentin 1983; Roth 1983),

hymenopterous parasitoids, and co rophagous Coleoptera (Roth

et al. 1991). Therefore more emphasis should be placed on

predators and parasitoids as biological control agents

(Legner 1978; Roth 1983).

Coprophagous flies. The adults and larvae of some

dipteran flies feed on dung, microorganisms (e.g. fungi,

bacteria) growing in the dung, or decomposing vegetable

matter (e.g., fungal spores and hyphae) (Merritt 1976).

Though some species of Muscidae ard Sarcophagidae may

compete with the horn fly in central Texas (Kunz 1978), the

populations of these flies are usually not as high as those

of coprophagous beetles, because they encounter predation

and competition. Small dung flies, such as Sepsidae, have

very high populations, but their biomass is considerably

less (Harris and Blume 1986). Mac ueen and Beirne (1975)

thought that dung-inhabiting Dipte a do not encounter food

limitations and Poorbaugh et al. (1968) regarded competition








34

for food or space as rare among ccprophagous Diptera.

In Australia, horn flies dev loping in cattle dung

experienced little competition for food or losses due to

predation from other arthropods (B5ornemissza 1960, 1970).

The effect of other coprophagous Ilies on the horn fly

population in North American pastures has not been reported.



Parasitoids

The Hymenoptera in dung, with the exception of ants,

are parasitoids. Blume (1985) listed 43 species of

parasitoid Hymenoptera associated with cattle dung. Twenty-

two of them, summarized by Fincher (1990), have been

reported as parasitoids of the horn fly in the United States

(Lindquist 1936; Thomas 1981; Thomas and Morgan 1972a; Figg

et al. 1983; Watts and Combs 1977; Combs and Hoelscher 1969;

Harris and Summerlin 1984; Schreiber and Campbell 1986).

Depner (1968) and Peck (1974) reported nine parasitoid

species from Canada. Although many species of parasitoids

have been reported to attack the horn fly, the most common

parasites are in the genus Spalanqia (Harris and Blume

1986).

Some staphylinid beetles have been recorded to

parasitoidize the pupae of horn flies. They are Aleochara

bimaculata Gravenhorst (Klimaszews i 1984), and Aleochara

sp. (Cervenka and Moon 1991). In these species the newly

hatched larva enters a fly puparium, develops within, and










emerges as an adult beetle. The adult beetles are

predacious on eggs and larvae of muscoid Diptera (Wingo et

al. 1974, Klimaszewski 1984). A report (Escher 1977) that

Tinotus planulus Notman and Tinotus sp. are parasitoids has

not been substantiated; and the report that Oxytelus incisus

is a parasitoid is certainly erroneous.

The level of parasitoidism of the horn flies can be as

high as 43% (Combs and Hoelscher 969) or 45% (Harris and

Summerlin 1984), but usually it i low and only occasionally

is the level adequate to reduce horn fly populations

significantly (Harris and Blume 1S86; Macqueen and Beirne

1975). Wharton (1979) collected thousands of horn fly pupae

from California in spring and summer, but no parasitoids

were found.

Ten hymenopterous species parasitoidal on the horn fly

pupae in Florida were reported by Escher (1977) and Butler

et al. (1981), with an overall level of parasitoidism of

10.5% (from 1.9 in April to 17.7 in August).



Ecology and Biology of Staphylinidae

Specific information on the biology, ecology and

behavior of many species of dung- nhabiting predatory

beetles, especially the Staphylin dae, is scant. However,

this information is very useful in evaluation of native and

exotic species as biological control agents of the horn fly,

in mass propagation of efficient species for initial release










and in establishment of programs in selected areas of the

country.

Hunter et al. (1991) reported 22 species of

Staphylinidae in their survey in Texas and described the

pattern of seasonal distribution and diel flight activity of

abundant species. Hanski and Hammond (1986) observed that

predatory species were more patchily distributed than

saprophagous species. Successes in rearing Staphylinidae

have been reported in North America (Mank 1923; Harris and

Oliver 1979; Hunter et al. 1986b) and in Europe (Hinton

1944; Paulian 1941) and Africa (Tawfik et al. 1976a, b, c).

Life history and habits have been observed of some

species of Philonthus (Mank 1923; Hunter et al. 1986a;

Tawfik 1976a,b,c), Platystethus (Hinton 1944; Legner and

Moore 1977) and Aleochara (Peschke and Fuldner 1977; White

and Legner 1966; Drea 1966). Dung-inhabiting Staphylinidae

have four stages: egg, larva, pupa and adult. There are

three larval instars for most subfamilies and two for

Paederinae (Frank 1991). The eggs are white, some with

sculptures in the surface of chorion, and hatch within 2-5

days. An egg burster was observed for the embryo within the

egg (Hinton 1944). The neonates of Aleochara gnaw a hole in

a fly pupa, enter and develop in the puparia (Peschke and

Fuldner 1977; White and Legner 1966; Drea 1966), and the

remaining genera are free living aid pupate in the dung or

soil under the dung pats (Hinton 1944; Mank 1923; Frank








37

1991; Hunter et al. 1986b, 1989; Tawfik et al. 1976a, b, c).

The immature stages have beer described for

Staphylininae (Frank 1991; Mank 1923; Hunter et al. 1989;

Tawfik 1976a, b, c; Hinton 1981a&;; Wu and Zhang 1990),

Oxytelinae (Frank 1991; Hinton 1944; Legner and Moore 1977),

Aleocharinae (Peschke and Fuldner 1977; White and Legner

1966), and other subfamilies (Frank 1991).














CHAPTER 3
THE ARTHROPOD COMMUNITY IN NORTH CENTRAL FLORIDA PASTURES


The presence of other insects in cattle dung has been

shown to reduce populations of the horn fly, Haematobia

irritans (L.) (Blume et al. 1970, Thomas and Morgan 1972b,

Macqueen and Beirne 1975). Hence, a basic study of the

insect fauna is essential for evaluation of regulation of

the horn fly and for consideration of further need of

introduction of biocontrol agents into an area. Several

comprehensive surveys have been conducted in North America

(Macqueen and Beirne 1975; Merritt and Sanders 1977; Mohr

1943; Sanders and Dobson 1966; Poorbaugh et al. 1968; Blume

1970; Valiela 1969b; Wingo et al. 1974), and over 400

species of arthropods have been collected in or on cattle

dung in the United States (Harris and Blume 1986). No

comprehensive study of the fauna cIf cattle dung has yet been

reported for Florida. Arthropod pecies that affect horn

flies vary geographically, therefore, biotic mortality of

horn fly immature stages in Florida may not be assessed

using the data of arthropod composition and abundance from

any other states. Results reported herein are a study on

the arthropod community in pastures in north central

Florida. The objectives of this tudy were to determine the










composition and abundance of the arthropod community

associated with cattle dung, and to monitor seasonal

distribution and diel activity of arthropods found.



Materials and Methods



A survey of the arthropod co munity associated with

cattle dung was conducted in July 1991 and from June 1992 to

December 1993 in pasture A and in September-December 1993

and 1994 in pasture B. Pasture A is 16 km (10 miles)

northeast of Gainesville, Alachua County, Florida, and

contained approximately 250 beef cattle at the time of the

study. Pasture B is on the south rn side of Gainesville and

contained 40 cattle at the time o the study. Two methods

of collection were used: pitfall traps baited with fresh

cattle dung, and emergence boxes (Fig. 3-1) that held entire

cowpats and trapped all emerging rthropods. A dilute soap

solution was added to the pitfall traps to drown trapped

arthropods and keep them clean and flexible for subsequent

processing. Ten to 15 pitfall traps were set twice each

month from May to October, and once each month from November

to April in pasture A; 20 traps were set in September and

October 1992 and 1993 in pasture B, respectively.

Arthropods captured in the traps iere collected after 24

hours and preserved in 70% alcohol for identification.

Between December 1992 and December 1993, cattle dung










pats about 24 hours old were collected from pasture A and

taken to the laboratory to extract the dung-associated

arthropods. Five dung pats were sampled twice a month from

May to October and once a month from November to April.

Dung pats were also sampled from pasture B in October 1992

and 1993. The pats were placed individually into emergence

boxes of a type used by G. T. Fincher (USDA-ARS, College

Station, Texas) who provided one, as a model for

construction of others. Each box was a gray plastic kitchen

box 46 cm long X 33 cm wide x 18 cm high. A 30 x 20 cm2

section was cut from each lid and replaced with a piece of

black cotton cloth to provide ventilation. A circular hole

(4 cm diameter) was cut through one end of the box. The lid

for a 7.5 cm high X 4 cm diam vial was perforated, then

glued and riveted to the box over the hole. When the vial

was screwed onto the box over the hole, it served as a

collection device to collect arthropods that attempted to

escape the emergence box by flight. In a similar manner, a

10 cm diam hole was cut through the bottom, and the lid of a

12.7 cm deep X 12.7 cm diam plastic cup was perforated, and

glued and rivetted over the hole nd then the cup was

screwed onto the lid. This device collected arthropods

walking and falling into it. The mouths of the vial and the

cup were each fitted with a hardware cloth funnel to prevent

insects from escaping back into the box. The vial and the

cup collected adult arthropods that left the dung in the























































Fig. 3-1. Emergence boxes used for extracting arthropods
from intact cowpats 24 hrs oid.









42

Table 3-1. Summary of invertebrates collected by two methods
in pastures in north-central Florida from 1991-1993.


Class and Order No. Families No. Species



Insecta
Orthoptera 2 3
Homoptera 3 5
Coleoptera 14 109
Diptera 20 35
Hemiptera 5 9
Lepidoptera 3 3
Hymenoptera 16 24
Dermaptera 1 1
Thysanoptera 1 1
Isoptera 1 1
Collembola 3 15

Arachnida
Acari 5 12
Araneae 1 4

Diplopoda 1 1

Nematomorpha 1 1
Nematoda 1 1

Mollusca 1 1


Total 226


Total


226










4
A Hydrophilidae
A Histeridae
U) S Carabidae
+





cn
2 -



F--
~I)


(/) Scarabaeidae 0 Staphylinidae
0) 45 -
a)
O60

z30 -



15 -

0

J F M A M J J A S 0 N D

Month 1993
Fig. 3-2. Seasonal distribution of five families of
Coleoptera most commonly collected by trapping (Mean+
Se). N = 10 X 2 for May to October and N = lo for the
remaining months.







































J F M A M Ji J A S 0 N D
Month 1993












Fig. 3-3. Seasonal changes in number of species of
Staphylinidae and Scarabaeidae in pastures of north
central Florida.










Table 3-2. Staphylinidae collected in pastures in north
central Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and (+++) abundant.


Trapped Extracted


Osorinae
Osorius sp. 1

Oxytelinae
Oxytelus incisus Motschulsky 617 2365
Platystethus spiculus Erichson 0 57
P. americanus Erichson 14 109
Anotylus nanus (Erichson) 65 79
A. insignitus (Gravenhorst) 129 67
Apocellus sphaericollis (Say) 15 26

Paederinae
Rugilus angularis (Erichson) 26 22
Achenomorphus corticinus 2 +
(Gravenhorst)
Lithocharis sororcula (Kraatz) 27 34
Thinocharis sp. 10 36
Homaeotarsus cinctus (Say) 2 +
Astenus fusciceps (Casey) 3 +
Scopaeus sp. 1 +

Staphylininae
Philonthus hepaticus Erichson 11 107
P. flavolimbatus Erichson 95 53
P. longicornis Stephens 91 38
P. rectangulus Sharp 2 +?
P. sericans (Gravenhorst) 5 26
P. ventralis (Gravenhorst) 16 36
Endius sp. 2
Gabronthus mgogoricus Tottenham 1
Platydracus tomentosus 4 0
(Gravenhorst)

Xantholininae
Phacophallus tricolor (Kraatz) 3 +
Lithocharodes ruficollis 8 +
(LeConte)
Neohypnus pusillus (Sachse) 15 119
N. attenuatus (Erichson) 6 32
N. emmesus (Gravenhorst) 2 11











Table 3-2. (continued)


Trapped Extracted


Aleocharinae
Acrotona hebeticornis Notman 336 59
Atheta sp. 207 93
Aleochara notula Erichson 507 ...
Gnypeta floridana Casey ? 2
Tinotus amplus Notman 59 179
T. brunnipes Notman 76 191
Thyasophila sp. 2
Falagria dissecta Erichson 2 +
Meronera venustula (Erichson) 3
Hoplandria sp. 1 1
sp. 2 1
Thecturota sp. 1
Alaobia scapularis (Sahlberg) 1
Athetini (genus?) 6

Tachyporinae
Bryoporus rufescens LeConte 4 +
Mycetoporus flavicollis 7 +
(LeConte)

Undetermined 265 5

* all Staphylinidae were determined by J. H. Frank.







47

12
0 Oxytelus incisus
9- Anotylus spp.

6

3
I.- O -0-



6 0 Aleochara notula
0
(U

= 2
4-



0 4Atheta sp.
zi
3 0 Tinotus spp.
Acrotona hebeticomis
2

1

0
I I I
J F M A M J J A S 0 N D
Month 1993


Fig. 3-4. Seasonal distribution of staphylinid species in
pasture A, collected by trapping. N = 10 X 2 for May
to October, and N = 10 for the remaining months.








48
emergence boxes. These arthropods either were in the adult

stage when the dung was collected, or had developed from

immature stages within the dung.

Insect diel activity was measured by pitfall traps

baited with fresh dung. Fifteen traps were run continuously

for 24 hours in pasture A in July and August 1992,

respectively. At each 6-hour interval, the bait in the

traps was replaced with fresh dung and the trapped insects

were collected.



Results and Discussion



Over the sampling period from 1991 to 1993, more than

50,000 invertebrates (Collembola were not included) were

collected in two pastures by two collecting methods. In

total, 226 species of invertebrates were collected and

identified. Arthropods found were 223 species belonging to

79 families in 14 orders. Coleoptera were ranked first in

number of species (109), Diptera second (35), and

Hymenoptera third (24) (Table 3-1).



Coleoptera

Staphylinidae held the most species (44) (Table 3-2),

followed by Scarabaeidae (27) and Carabidae (20). Four

species each of Histeridae and Hydrophilidae were collected

(Table 3-4). Staphylinidae accounted for the greatest








49
number of individuals, followed by Scarabaeidae. The numbers

of Hydrophilidae, Carabidae and Histeridae were much lower.

Staphylinidae. An estimated il0,000 adult Staphylinidae

were collected and over 7000 were identified to species

level. The specimens belong to seven subfamilies, of which

Staphylininae, Aleocharinae, Oxytelinae and Paederinae were

most abundant (Table 3-2), Xantholininae were common, and

Tachyporinae and Osorinae were rare. Oxytelus incisus and

Aleochara notula were abundant from trapping and extracting,

Tinotus spp. were abundant in extractions, and Acrotona

hebeticornis and Atheta sp. were abundant in pitfall traps.

Anotylus spp., Philonthus flavolimbatus, P. longicornis and

Neohypnus pusillus were common from trapping or extracting.

Platystethus americanus, Philonthuls hepaticus were common in

extraction. Platystethus spiculus were only collected by

extracting and Platydracus tomentosus were only collected by

trapping.

Members of the genus Philonthus have been well

documented to be predators of fly immature stages (Hammer

1941; Harris and Oliver 1979; Hunter et al. 1989; Macqueen

and Beirne 1975; Roth 1982; Thomas and Morgan 1972b). Six

Philonthus species were collected in this study. P.

hepaticus, P. flavolimbatus, and P1. longicornis were more

common than the other three species, of which P. rectangulus

was rare. The adventive species Pi. longicornis and P.

flavolimbatus were common and have been reported to be










efficient predators of the horn fly (Roth 1982; Harris and

Oliver 1979; Fincher and Summerlin 1994). Except for P.

hepaticus, local species (P. ventralis, P. sericans and P.

rectangulus) were less commonly collected than the two

adventive ones. This indicates that these adventive species

are well established and competed successfully with native

Philonthus species.

Aleochara notula was very abundant during the survey.

The adults prey on fly eggs and larvae of cyclorrhaphous

Diptera, and its immature stages are solitary

ectoparasitoids of fly pupae within the host puparia

(Klimaszewski and Blume 1986; Harris and Blume 1986). They

were collected from dung in emergence boxes from the time

the dung was placed in the boxes until about 3 weeks

thereafter. A sarcophagid [Ravini:a delicta (Walker)] was

found to be the main host of this beetle in the dung,

followed by Brontaea debilis (Williston) and B. cilifera

(Malloch) [Muscidae]. No horn fly pupae were found to be

parasitoidized by Aleochara notulal. Twenty-three A. notula

were reared from Ravinia delicta pupae in one cowpat in

August 1993. The adults of A. notula were found to be

abundant and active in fresh to one week old dung.

Oxytelus incisus and Atheta sip. were abundant in the

survey. They may play an important role in reducing horn

fly populations in north central Florida if they prey on

horn fly immatures. Valiela (1974') cited Oxytelus










tetracarinatus Block and Atheta sp. as small predators but

did not present evidence. It is reported that Platystethus

spiculus (Palomino and Dale 1989), P. americanus (Mohr 1943;

Cervenka and Moon 1991) and Falagria dissecta (Valiela 1974)

are predators of flies. These species, however, were not

abundant in this study.

Staphylinidae were collected year round. Numbers were

low during the winter, built up slowly in the spring, and

peaked in the late summer or fall. The seasonal abundance

for 1993 showed bimodal peaks in July-August and November

(Fig. 3-2). The first peak was mainly composed of Oxytelus

incisus, and Aleochara notula (Fig. 3-4); the second peak

was mainly composed of Anotylus spp., Tinotus spp. and

Acrotona hebeticornis (Fig. 3-4).

The number of species of Stap hylinidae was low in the

spring, increased in the early summer, peaked in June-August

(14-16 species), and then declined, (Fig. 3-3). Different

staphylinid species were collected, during the different

seasons of a year. Anotylus insignitus and A. nanus were

mainly collected during the winter and spring. Meronera

venustula and Mycetoporus flavicollis were mainly collected

in the spring. Tinotus brunnipes, T. amplus and Acrotona

hebeticornis were mainly collected during the winter.

Atheta sp. were collected each month. Philonthus spp., A.

notula, and 0. incisus were mainly collected during the

summer and fall. Most of the remaining species were










collected sporadically.

Diel activity was also different among the species. A.

notula and Philonthus spp. were mainly collected during the

day time, but Tinotus spp., Acrotona hebeticornis, Atheta

sp. and Oxytelus incisus were mainly collected during the

night.

Adult Staphylinidae began to leave the dung placed in

the emergence boxes a few hours after the dung was brought

into the laboratory. The number increased the next day, was

most abundant within about a week, and then declined. Most

of the staphylinid beetles left the boxes within two weeks

after dung was collected and placed in them. Those

extracted later developed from the immature stages present

in the dung when it was collected.

Successful biological control of dung-inhabiting pest

flies depends on ecological factors, especially seasonal

distribution and the habitat preference of natural enemies.

In north central Florida, horn flies are active all year

round, without overwintering (Wilkerson 1974), and become

most active from May to October. Staphylinidae are the most

important predators of the horn fly (Fincher 1990; Harris

and Blume 1986) elsewhere, but our survey results showed

that the numbers of Staphylinidae did not reach their peak

until July, especially Philonthus spp. and A. notula (Fig.

3-4) which have biocontrol potential for the horn fly.

Because of this 'lag' time, numbers of horn flies increase










Table 3-3. Scarabaeidae collected in pastures in north
central Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and (+++) abundant.


Trapped Extracted



Scarabaeinae
Onthophagus gazella F. 971 ...
0. taurus (Schreber) 71 ++
0. pennsylvanicus Harold 300 ...
0. hecate blatchleyi Brown 259 ...
0. striatulus floridanus 1
(Beauvois)
0. oklahomensis Brown 165 ++
0. tuberculifrons Harold 286 ...
Copris minutus (Drury) 7 +
Canthon pilularius (L.) 42 ++
Boreocanthon probus (Germar) 1
Phanaeus vindex MacLeay 31 +
P. igneus floridanus d'Olsoufieff 4

Cetoniinae
Euphoria sepulchralis (F.) 4

Hybosorinae
Hybosorus illigeri Reiche 4

Aphodiinae
Ataenius gracilis (Melsheimer) 9
A. picinus Harold 55 ++
A. fattigi Cartwright 2
A. imbricatus (Melsheimer) 39 +
A. platensis Blanchard 1
Aphodius campestris Blatchley 15 +
A. lividus (Olivier) 1077 ...
A. parcus Horn 15 +
A. fimetarius (L.) 22 +
A. cuniculus Chevrolat 8

Geotrupinae
Geotrupes egeriei Germar 4
Bradycinetulus ferrugineus
(Beauvois) 2
Dynastinae
Strategus splendens (Beauvois) 1

* Scarabaeidae were determined by M. Thomas, P. Skelley, P.
Choate and J. McNamara.


























































Fig. 3-5. Seasonal distribution of scarabaeid species in
pasture A, collected by trapping. N = 10 X 2 for May to
October and N = 10 for the remaining months.










rapidly in the spring before populations of Staphylinidae

increase accordingly. If predator abundance could be

increased and maintained at a high level during this period

by inundative releases of native or introduced species with

similar activity patterns, horn fly numbers might be

reduced.

Scarabaeidae. Over 8,000 scarab beetles were collected

and more than 5,000 were identified to species (Table 3-3).

They belong to 27 species of six subfamilies, of which

Scarabaeinae (12 species) and Aphodiinae (10 species) were

predominant. Onthophagus Qazella, and Aphodius lividus

were the most abundant (Table 3-3), followed by 0.

pennsylvanicus, 0. h. blatchlevi, 0. oklahomensis and 0.

tuberculifrons. 0. taurus, Canthon pilularius, Phanaeus

vindex, and Ataenius picinus were commonly collected.

Unlike Staphylinidae, very few Scarabaeidae were

collected from January to March. Their numbers started to

increase in April, the first peak was in May, and the second

and third in July and November 1993, respectively (Fig. 3-

2). The peak in May was mainly contributed by A. lividus;

the one in July by 0. gazella and supplemented by A.

lividus; and the November peak by 0. pennsylvanicus, o. h.

blatchleyi, 0. oklahomensis, and 0. tuberculifrons (Fig. 3-

5).

High numbers of A. lividus in the early summer (May

1993) contributed to the breaking of the dung pats. These










small beetles swarmed into the dung pat within a few house

after the dung was deposited. One day later the dung pat

was mostly consumed and totally broken up, and then became

dry in 4-5 days. Flies and other immature insects were not

found abundant during this time. 0. gazella, which were

released in Texas in 1972 and found in Florida in 1975

(Fincher 1990), were abundant only in July and August. It

was reported that this beetle remo ed over 80% of dung from

the soil surface during its peak activity in Texas (Harris

1981). A similar situation was observed but not measured in

July and August in the present survey. 0. taurus, too, is

not a native species (Fincher 1990), but its numbers were

much lower than those of 0. gazelle and the other native

Onthophaqus species.

The remaining five species of Onthophagus, which were

abundant during the fall and early winter (Fig. 3-5), were

less important than A. lividus and 0. gazella in

decomposition of dung.

No scarab beetles were collected in January, but the

number of species increased from February, was highest in

June to August (14-15 species), th n declined. There was a

second peak in November (Fig. 3-3).

Most 0. gazella, other Onthophagus spp. and A. lividus

were collected during the daytime.

Though small numbers of AtaenLus spp. and Aphodius spp.

were extracted 1-3 weeks after the pats were collected and








57

brought into the laboratory, most Scarabaeidae left the dung

placed in the emergence boxes within two days.

The results showed that the numbers of scarab beetles

built up later in the year than the numbers of horn flies.

Introduction of more efficient coleopteran competitors or

inundative releases of native species in the spring may

reduce the horn fly numbers in north central Florida.

Carabidae. A total of 20 species of carabids was

collected, but none were abundant. However, Tachys sp.,

Ardistomis viridis, Stenocrepis cauatuordecimstriata and

TetraQonoderus intersectus were common (Table 3-4). Some

large-sized species did not occur in the dung at all, such

as Calosoma savi and Pasimachus sp ., even though Pasimachus

spp. have been reported to be associated with bovine

droppings (Blume 1985). Some smal and medium-sized

species were commonly extracted fr m the dung. Since they

are general predators, they may be important in reducing

horn fly populations. Mohr (1943) and Hammer (1941)

reported, respectively that larvae of Carabidae were

predators of fly larvae in dung. 9nly two Galerita larvae

were collected in the dung, but may adult Carabidae

extracted developed from immature stages in the dung.

Most species in this family w~re collected

sporadically. The number of individuals was low all the

year and there were no obvious pea s (Fig. 3-2). Tachys

sp., and S. quatuordecimstriata were collected in most










Table 3-4. Coleoptera collected in pastures in north central
Florida, with exclusion of St phylinidae and
Scarabaeidae. The occurrence codes for species not
enumerated: (+) rare, (++) co mon, and (+++) abundant.


Tr pped Extracted



Carabidae
Tachys sp. 36 ++
Panagaeus fasciatus (Say) 1
Evarthrus morio Dejean 3
Calosoma sayi Dejean 6
Ardistomis viridis Say 22 +
A. puncticollis Putzeys 9 +
Dyschirius sp. 2
Clivina sp. 2
Aspidoglossa subangulata Chaudoir 7
Stenolophus infascatus (Dejean) 1
Selenophorus paliatus F. 9 +
S. discopunctatus Putzeys 2
S. fossulatus Dejean 4
Amblygnathus irepennis (Say) 1
Amara sp. 1
Stenocrepis quatuordecimstriata 36 ++
Chaudoir
Galerita sp. 1
Tetragonoderus intersectus 20 ++
Haldeman
Pasimachus sublaevis Dejean 8
P. marginatus (F.) 2

Histeridae
Hister coenosus Erichson 26 11
Phelister haemorrhous Marseul 5 2
Acritus ignobilis Lewis 22 7
Saprinus pennsylvanicus (Paykull) 28 6

Hydrophilidae 85
Sphaeridium lunatum (L.) 34 5
Cryptopleurum subtile Sharp 3
Cercyon variegatus Sharp 199 42
C. atricapillus (Marsham) 134 31

Tenebrionidae
Gondwanocrypticus obsoletus(Say) 13 4
Poecilocrypticus formicophilus 2 4
Gebien










Table 3-4 (continued)


Trapped Extracted


Cicindelidae
Cicindela puntulata Olivier 4
C. scutellaris Dejean 1
Megacephala virginica L. 1

Scolytidae (1 sp.) 27 ++

Ptiliidae (1 sp.) 32 ++

Elateridae (3 spp.) 13

Anthicidae
Anthicus sp. 6 +

* Carabidae were determined by P. Choate and M. Thomas;
Histeridae by P. Kovarik; Hydrophilidae by A. Smetana; and
Cicindelidae by P. Choate.










months of the year. A. viridis were collected in the

summer, and T. intersectus and P. sublaevis were collected

during the summer and fall.

Most species in this family were collected during the

day time.

Hydrophilidae. Sixteen species of Hydrophilidae have

been reported associated with cow dung in the continental

USA (Fincher 1990), but only four species were collected in

the present survey. Sphaeridium scarabaeoides (L.) has been

considered to be an important predator of horn flies (Bourne

and Hays 1968; Hammer 1941; Macqueen and Beirne 1975; Mohr

1943; Poorbaugh 1966; Sanders and Dobson 1966). S. lunatum

were commonly collected. It is similar in size to S.

scarabaeoides and these two specie may have similar

predation potential against the horn fly. Cercyon

varieqatus and C. atricapillus were abundant (Table 3-4)

but they have not been considered -o be predators (Sanders

and Dobson 1966; Hafez 1939; Merritt 1976; Thomas and Morgan

1972b).

More Hydrophilidae were collected in the spring and

winter than in the summer and fall (Fig. 3-2).

Histeridae. Twenty-two species of Histeridae have been

reported from dung in the continental USA (Fincher 1990).

Only four species of these beetles were collected in this

survey. The most abundant was Hister coenosus, which is

medium-sized and has been considered to be the most








61

efficient histerid in reducing hor fly populations

(Summerlin et al. 1982, 1984c, 199 ). The other three

species were less common, of which Saprinus pennsvlvanicus

and Phelister haemorrhous have alsD been reported to be

predators of horn fly immatures (Summerlin et al. 1982,

1991).



Diptera

In total, 35 species of Diptera belonging to 19

families were collected (Table 3-5). Abundant species of

higher dipterans included Haematobia irritans, Brontaea

debilis, B. cilifera and Neomyia cornicina in the family

Muscidae; and Ravinia derelicta, R. floridensis, Helicobia

morionella and Oxysarcodexia ventricosa in the family

Sarcophagidae. Abundant lower dun -inhabiting dipterans

included Coproica sp. (Sphaeroceri ae), Palaeosepsis

insularis (Sepsidae), Aphodiplosis triangularis and

Lestodiplosis sp. (Cecidomyiidae), Bradysia coprophili

(Sciaridae) and Sylvicola notialis (Anisopodidae).

H. irritans arrives earliest at the fresh dung and

emerges earliest of all the flies from the dung. Previous

study shows that this fly was active all year round in north

central Florida (Wilkerson 1974). The fly numbers were

observed to be low during the earl spring, became high in

May and peaked in August to September on the cattle in 1993.

The adults were extracted from dung pats collected in










Table 3-5. Diptera collected in pastures in north central
Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and (+++) abundant.


Trapped Extracted


Muscidae
Haematobia irritans L. 0 226
Brontaea debilis (Williston) ++ 375
B. cilifera (Malloch) ++ 198
Pseliphephila sp + +
Neomyia cornicina (F.) ++ 497

Sphaeroceridae
Coproica sp. + + 2221

Sepsidae
Palaeosepsis insularis (Williston +++ 1358

Ephydridae
Discocerina obscurella (Fallen) 2

Sarcophagidae
Ravinia derelicta (Walker) ++ 226
R. floridensis (Aldrich) ++ 79
Udamopyga niagrana (Parker) 2 +
Oxysarcodexia ventricosa (Wulp) ++ 38
Helicobia morionella (Aldrich) ++ 33

Cecidomyiidae
Aphodiplosis triangularis (Felt) ++ 1142
Lestodiplosis sp. (new) +++ 760
Neolasioptera sp. ++ 620
Aprionus sp. ++ 18

Bibionidae
Plecia nearctica Hardy + +

Dolichopodidae
Sciapces sp. ++ +
Syntormor sp. 35
Sp. 3 ++ +

Empididae
Drapetis vanthopoda Will. ++ 47

Milichiidae (1 species) 4 ++








63

Table 3-5 (continued)


Trapped Extracted


Ceratopogonidae
Forcipomyia brevipennis (Macquart ++ 63

Scatopsidae (1 species) + 42

Tachinidae (2 species) + 13

Psychodidae (2 species) + 129

Sciaridae
Bradysia coprophili (Lintner) ++ 107

Phoridae
Megaselia sp. + 16

Anisopodidae
Sylvicola notialis Stone ++ 266

Chloropidae
Monochaetoscinella nigriconis (Loew) ++ 32

Drosophilidae (1 species) ++ 44

Tabanidae (1 species) + 4

Tipulidae (1 species) 5 4

Muscidae were determined by D. Deonier, G. Gagn6, and G.
Steck; Sarcophagidae by G. Steck and G. Dahlem; Bibionidae,
Sepsidae and Sphaeroceridae by G. teck; Cecidomyiidae and
Sciaridae by J. Gagn6 and W. Groga ; Phoridae by D. Deonier;
Ceratopogonidae by W. Grogan and G Steck; Dolichopodidae by
K. Ahlmark; Empididae and Chloropidae by G. Steck; and
Anisopodidae and Ephydridae by W. Wirth.








64

pastures year round and were abundant from July to October.

Brontaea spp. were collected in the late summer and fall,all

year round and were very abundant uring the summer and

fall. Ceratopogonidae, and Dolicho odidae were mainly

collected in the fall. Psychodidae and Anisopodidae were

mainly collected in the late-fall nd winter. Bibilionidae

were collected only in July 1991 and in July-August 1992.

Larvae of Ravinia lherminieri (Robineau-Desvoidy) have

been reported to be predators of other dung-inhabiting flies

(Pickens 1981). R. derelicta was bundant and might play

the same role as R. lherminieri in reducing horn flies.

Sphaeroceridae, Sepsidae Scatopsid e, and Psychodidae are

primarily dung feeders (Skidmore 1991; Valiela 1974).

Sphaerocerid flies are reportedly horetic on scarab beetles

in Florida (Sivinski 1983). Cecidomyiidae and Sciaridae are

primary fungus feeders, but the larvae of Lestodiplosis sp.

are predacious (Gagnd, pers. comm. The effect of these

larvae on horn fly and other fly pc pulations has not been

measured. Empididae and Dolichopodidae have also been

mentioned as predators (Hammer 1941), but their effect on

horn and other flies in the dung is unknown. The larvae of

Bibionidae are big and active in the dung and seem to be

predators of flies, but they are r ported to invade the dung

from surrounding soil (Skidmore 19 1). Chloropidae,

Ephydridae, Drosophilidae, Tipulidae and Tabanidae occur

casually in cow-dung. According to Skidmore (1991), the










larvae of most chloropid species develop in the stems of

grasses; the larvae of Ephydridae evelop in a range of

materials, including muddy soil, rDtting vegetation, or cow-

dung; and Drosophilidae develop mostly in decaying plant or

animal matter in which fermentation is taking place.

Drosophilidae were abundantly seen in decaying mushrooms

growing in pastures in the present study. Tipulid larvae

occur in dung, but belong to the soil fauna.



Hymenoptera

A total of 24 species belonging to 15 families of

Hymenoptera was collected. The family with the most

represented species was Formicidae, of which the red

imported fire ant (RIFA), Solenops s invicta, was abundant

all year round and peaked in September-October. RIFA has

been well documented as a predator of flies (Laurence 1954;

Bruce 1964), in particular, of horn flies (Howard and Oliver

1978; Summerlin et al. 1984b; Schmidt 1984). The effect of

RIFA on the horn fly will be discussed in chapter 5.

Two species of eulophids were collected: Aprostocetus

spp. have been recorded to be para, itoids of gall-midges,

and Trichospilus spp. as parasitoids of fly puparia

(Krombein et al. 1979). Among the Braconidae collected, one

species (Aphidiinae) is a parasitold of aphids. The other,

Aphaereta pallipes, was commonly co elected by trapping and

rearing from extracted sarcophagid puparia; its hosts










Table 3-6. Hymenoptera collected in pastures in north
central Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and (+++) abundant.


Trapped Extracted


Pompilidae
Anoplius sp. +
sp. 2 (Genus?) + +
Eulophidae
Trichospilus sp. 5 +
Aprostocetus sp. 2 ++
Braconidae
Aphaereta pallipes (Say) 17 ++
Sp. 2 (Aphidiinae) + +
Pteromalidae
Spalangia cameroni Perkins 10 ++
Formicidae
Solenopsis invicta Buren 38250 ...
Pheidole dentata Mayr + +
P. metallescens Emery + +
Cyphomyrmex sp. + +
Pseudomyrmex mexicana + +
Hypoponera opaciceps + +
Ichneumonidae
Gambrus ultimus (Cressson) 3 +
Mymaridae
Gonatocerus sp. + +
Eucoilidae
Trybliographa sp. 12 ++
Kleidotoma sp. + +
Rhoptromeris sp. + +
Diapriidae
Trichopria sp. + +
Encyrtidae
Adelencyrtus odonaspidis + +
(Fullaway)
Scelionidae
Telenomus sp + +
Sphecidae
Crabro rufibasis (Banks) + +
Platygastridae (1 species) + +
Dryinidae
Gonatopus sp. + +

* Braconidae, and Pteromalidae wer determined by V. Gupta;
Diapriidae and Scelionidae by P. Marsh; Pompilidae,
Sphecidae, Dryinidae, and Eucoilidae by A. Menke; Eulophidae
and Mymaridae by M. Shaufer; and Formicidae by D. Williams.









include house flies, flesh flies and horn flies (Wharton

1977; Sanders and Dobson 1966). Three species of Eucoilidae

Trybliographa sp., Kleodotoma sp., and Rhoptromeris sp.)

were represented; three genera have all been recorded as

parasitoides of fly puparia (Krombein et al. 1979).

Gonatocerus spp. are recorded as parasitoids of homopteran

eggs (Krombein et al. 1979). SpalanQia cameroni

(Pteromalidae) was reared out from the puparia of H.

irritans and Brontaea cilifera; it has been recorded as a

parasitoid of horn fly pupae by Es:her (1977) and Butler et

al. (1981) in Florida. Trichopria spp. are recorded as

parasitoids of fly puparia (KrombeLn et al. 1979).

Telenomus spp. are recorded as parasitoids of Hemiptera and

Lepidoptera (Krombein et al. 1979) Gambrus ultimus is

recorded as a parasitoid of lepidopterous larvae and has

been found in lepidopterous cocoons (Townes and Townes

1962).



Other Insects and Animals

Orthoptera. A cricket, NemobIus fasciatus (DeGeer),

has been reported to prey on horn fly pupae under laboratory

conditions (Bourne and Nielsson 1967). Crickets were

commonly collected in pastures in the present study and in

previous studies (Bourne and Nielsson 1967; Schoenly 1983).

Acari. Mites were commonly collected by trapping and

I










Table 3-7. Miscellaneous invertebrates collected in pastures
in north central Florida. The occurrence codes for
species not enumerated: (+) r re, (++) common, and
(+++) abundant.


Trapped Extracted


Othoptera
Gryllidae
Gryllus rubens Scudder ++ +
G. firmus Scudder ++ +
Gryllotalpidae
Scapteriscus vicinus Scudder + +

Homoptera 28
Cercopidae
Prosapia bicincta (Say) + +
Cicadellidae
Exitianus exitiosus (Uhler) + +
Planicephalus flavicosta (Stal) + +
Delphacidae
Toya andromeda (Van Duzee) + +
T. propinqua (Fieber) +

Hemiptera 162 31
Nabidae (1 species) +
Cydnidae
Pangaeus bilineatus (Say) ++ +
Reduviidae
Xylocoris galactinus Reuter ++ +
Anthocoridae
Lasiochilus pallidulus Reuter ++ +
Orius sp. +
Lygaeidae
Pachybrachius basalis (Dallas) +
Paromius longulus (Dullas) + +
Geocoris uliginesis (Say) + +
Blissus insularis Barber + +

Lepidoptera
Tineidae (1 species) 1
Pyralidae
Herpetogramma
phaeopteralis (Guen~e) 8
Noctuidae
Spodoptera frugiperda (Smith) 42 ++










Table 3-7 (continued)


Trapped Extracted


Collembola
Hypogastruridae
Hypogastrura sp. ... ...
Xenylla welchi Folsom ... ...
Neanura muscorum Templeton ... ...
Sminthuridae
Sminthurus jupiterensis Snider (new) ++ ...
Bourletiella gibbonsi Snider ++
Entomobryidae
Lepidocyrtus violaceus Fourcroy ++ ...
L. cf. helenae Snider ++ ++
L. cyaneus (Tullberg) ++ ...
Tomocerus elongatus Maynard ++ ++
Salina wolcotti Folsom ++ ++
Salina (sp. new) + ++
Seira brasiliana (Arl6) + ++
S. brasiliensis Snider ++ ++
S. caheni Jacquemart ++ ++
Entomobryoides sp. + +

Dermaptera
Labiduridae
Labidura riparia (Pallas) 8 +

Thysanoptera (1 species) + +

Isoptera (1 species) 1 +

Arachnida (Acari) 675 ...

Anistidae
Prostigmata sp. + +
Parasitidae
Parasitus fimetorus (Berlese) ++
Parasitus sp. + +
Macrochelidae
Glyptholaspis confusa (Foa) ++ ++
Glyptholaspis sp. + +
G. pamericana Berlese + +
G. fimicula (Sellnick) + +
Microcheles mammifer Berlese + +
Microcheles sp. +
M. peniculatus Berlese ++ ++
M. muscaedomesticae (Scopoli) + +
Oribatidae (1 species) 187 ++











Table 3-7 (continued)


Trapped Extracted


Arachnida (Araneae) 134 ++
Linyphiidae
Eperigone banksi Ivie & Barrows +
Erigone autumnalis Emerton +
Meinoneta sp. +

Diplopoda
Polyzoniida (1 species) ++ +

Nematomorpha
Gordius sp. 3

Nematoda
Rhabiditidae
Coarctadera coarctata (Leuckart) ++

Mollusca 3


* Acari (mites) were determined by G. Krantz and M.
Hennessey; Araneae (spiders) by G. Edwards; Collembola by R.
Snider; Orthoptera, Hemiptera and Homoptera by F. Mead and
T. Henry; Lepidoptera by D. Habeck; Nematodes by K. Nguyen
and R. Esser; and Dermaptera by J. Watts.










by insect colonizers, including scarab and staphylinid

beetles, and muscid, sarcophagid, sphaerocerid and sepsid

flies. These mites were frequently found on the dung surface

in the field. Four families of mites, Anistidae (one

species), Parasitidae, Macrochelidae, and Oribatidae, were

collected, among which members of Parasitidae and

Macrochelidae have been reported to be predators of fly

immatures (Valiela 1974; Krantz 1983; Halliday and Holm

1987; Anderson 1983). Macrochelids have been considered to

have biological control potential of house, bush, face and

horn flies (Axtell 1963; Krantz 1983; Halliday and Holm

1987; Anderson 1983), in particular, M. muscaedomesticae

(Axtell 1963) and M. peniculatus (Roth et al. 1988b;

Halliday and Holm 1987). Oribatids occur in the old dung and

are thought to feed on the dung or on associated fungi

(Skidmore 1991).

Araneae. Spiders were often found in dung. They are

general predators, preying on whatever they meet and are

able to handle.

Millipedes. Millipedes are associated with decaying

plant materials, upon which they feed. They were commonly

collected by trapping and extracting and were abundant in

the fall and winter. There are no data on the relationship

between millipedes and flies.

Collembola. Springtails were abundantly collected by

trapping and extracting. They feed on decaying plant










materials and fungi and are often numerous in older dung.

They mostly belong to the soil fauna (Skidmore 1991). They

also provide food for small insect predators.

Hemiptera. Hemiptera were collected frequently by

trapping and occasionally by extraction. Many species in

this order are considered to be casual visitors to dung.

Species in the family Lygaeidae have been found to prey on

other insects in the dung (Skidmore 1991).

Hematomorpha and Nematoda. Gordius sp. (horse hair

worm) is a parasite of Coleoptera Dermaptera, Diptera,

Hemiptera, and Hymenoptera. It was recorded from the German

cockroach and Florida wood cockroach in Florida (Esser

1980). Coarctadera coarctata parasitizes staphylinids,

hydrophilids, and scarabs (Frank 1982).

The remaining groups of miscellaneous arthropods and

other animals collected have been recorded infrequently in

the past, and their interactions with flies are poorly

known.














CHAPTER 4
EFFECT OF ARTHROPOD PREDATORS ON HORN FLY SURVIVORSHIP
IN PASTURES AND UNDER LABdRATORY CONDITIONS




The results of an arthropod community survey in north

central Florida (chapter 3) showed that over 220 species of

invertebrates are present in association with dung. The

dung arthropod community has been shown to reduce horn fly

populations in the USA (Blume et al. 1970; Thomas and Morgan

1972b; Kunz et al. 1972; Roth 1989), Canada (Macqueen and

Beirne 1975) and Australia (Fay et al. 1986, 1990). Insect

predators are reported as the primary biotic mortality

factor of the horn fly (Thomas and Morgan 1972b; Macqueen

and Beirne 1975; Harris and Blume 1986; Roth 1983, 1989).

Studies have shown that the most important predators are

coleopterans of the families Staphylinidae, Hydrophilidae,

and Histeridae (Bornemissza 1968; Fay 1986; Legner 1986;

Roth 1989; Sanders and Dobson 1969; Thomas and Morgan

1972b). Philonthus beetles (Staphylinidae) have been shown

to have high potential for predation on horn fly immatures

(Roth 1982; Fincher and Summerlin 1994; Harris and Oliver

1979). In Florida, only Escher (1677) and Butler et al.

(1981) reported parasitism of horniflies, but there have

been no studies of the effect of the whole arthropod

73










community, especially predators, on horn flies.

Objectives of the following study were to determine

horn fly mortality caused by the whole arthropod community

in pastures, and to evaluate the predation potential of

individual predatory species on horn flies.



Materials and Methods



Tests of the Arthropod Community on Horn Fly Survivorship

Laboratory studies on horn fly survivorship

Survivorship of horn flies was studied in the

laboratory before initiating field studies. The horn flies

used were from the colony maintained by Dr. J. F. Butler,

Department of Entomology/Nematology, University of Florida.

Standard colony rearing methods were developed by Greer

(1975) and modified by Okine and Butler (1995). Larval

medium was the mixture of frozen and thawed manure and

peanut hulls. The adults were fed with bovine blood. The

colony was maintained in an environmental chamber at 27

3C and 75 5% RH with continuous light.

Manure used for experiments was collected in a pasture

within 30 min after its deposition, frozen for 48 hours and

thawed for 24 hours. The tests were conducted at 270C, 50-

60% RH and 14:10 (L:D) photoperiod.

Artificial pats were prepared by following the








75

procedure of Thomas and Morgan (1972b). Horn fly eggs less

than 4 hours old were removed from the egg collection tray,

suspended in well water and pipetted on paper towel strips

moistened with well water. Twenty-five eggs were counted on

each towel strip. A metal hoop, 20.32 cm diameter and 5.08

cm high, was placed on a section of grass in a large metal

pan. The area within the hoop was moistened with well

water. One-hundred eggs [25 eggs on each of four paper

strips (6 cm long X 2 cm wide)] were placed on the grass

within the hoop, each strip being placed at each of four

side of the edge. The area within the hoop was covered with

manure, the hoop was removed and a simulated manure pat was

formed (approximately 25 cm diam X 5 cm high). The

simulated manure pats were held in a rearing room for 7-8

days and then covered with cone traps to collect emerged

horn flies. Six replicates were conducted.



Field studies on horn fly mortality

Field mortality of the horn fly was evaluated during

July and August 1992 at pasture A where an arthropod survey

was conducted at the same time (chapter 3). Horn fly eggs

from the laboratory colony were used. Manure collection and

artificial pat formation were performed according to the

procedure for laboratory survival studies.

Two trials were conducted, one each in July and August.

Twenty simulated manure pats for e ch trial were formed










along the outside of the fence surrounding plots in the

pastures. Adjacent pats were separated by 10 m. Egg-seeding

procedures (100 eggs/pat) followed those for laboratory

survival studies. Ten pats (odd numbers) were at first open

to allow other insects to come to the dung and then covered

by cone traps on the 8th day after egg-seeding; the other 10

(even numbers) were covered immediately after egg-seeding.

The cone traps (Fig. 4-1) used for covering the seeded

cowpats were constructed of a wire frame and wrapped with

fine saran screen. The trap was 30.5 cm diameter (bottom)

and 50.8 cm high. A circular hole (5 cm diameter) was cut

through the screen on the top of the trap, and a 8.89 cm

high X 5.08 cm diameter vial was screwed on (the lid of the

vial was perforated, then glued and rivetted to the trap) to

cover the hole to collect horn flies that emerged from the

pat. The mouth of the vial was fitted with a saran screen

funnel to prevent insects from esclping back to the trap.

The vials were checked daily from the 8th day after egg-

seeding until two days after the list horn flies were found.

Differences in mean numbers of horn flies emerged

between treatments were analyzed by a Student's t-test

(SigmaPlot 1994). Before the t-test was conducted, the

numbers of the horn flies were transformed by log (N+1) to

satisfy equal variance and normality assumpted by t-test.










Determination of Predation Rates of Predators

Collection and colonization of predators

Adults of predatory beetles ware extracted using the

emergence cages (Fig. 3-1) from the pats collected in

pastures A and B. Colonies of predatory beetles were

initiated and maintained by following the method of Hunter

et al. (1986b) for Philonthus. Five to 10 pairs of beetles

were placed in 5 cm high X 15.24 c diameter plastic

containers with several moist paper towels crumpled one on

top of the other. Two circular holes (2.5 cm diam) were cut

through the lid; one was covered by fine cloth screen to

allow for a free flow of air; the other was open for

insertion of predators and prey (horn fly eggs). Horn fly

eggs (500-1000) suspended in distilled water were pipetted

on the surface of the towels twice a week as a food source.



Predation rate test under laboratory conditions

For observation of predation, predators were confined

individually in Petri dishes (5.5 cm diameter X 1.3 cm

high), lined on the bottom with a moist paper towel. A

water-soaked cotton ball was provided for humidity, and

horn fly eggs and 1st instar larva were provided for food.

An observation cage as described by Hinton (1944) was also

made for rearing P. americanus in the laboratory. The cage

consisted of a well cut in a piece of styrofoam. The well








78
was covered by a piece of microscope slide. Oviposition and

development of the predators at different stages were

recorded daily.

Predators used in this study included 22 staphylinid

species, two hydrophilid species, one histerid species, two

carabid species, one anthicid species, and one tenebrionid

species.



Predation rate test under simulated field conditions

Predation tests for Philonthus species were conducted

in testing cages (Fig. 4-2), which are made of 15.24 cm

diameter X 5 cm high plastic containers. Moist, sandy loam

soil was placed in each container (1 cm deep). Horn fly

eggs (100) suspended in well water were pipetted on a piece

of doughnut-shaped paper towel (13 cm diam), with a circular

hole (9 cm diam) cut in the center. The paper towel was

placed on the loam soil and covere( with a 100 g cattle

dung. This arrangement allowed the fly eggs to be covered

by the edge of the artificial pat, thereby simulating

natural horn fly oviposition behavior (McLintock and Depner

1954). Immediately after the eggs were seeded, one or two

adult Philonthus (mixed sexes) of each species were added to

the pat through a circular hole (2.5 cm diameter) on the

lid. Another circular hole was cut through the lid of the

container and an 8.89 cm high X 5. 8 cm diameter vial was

screwed on (the lid of the vial was perforated and rivetted
























































Fig. 4-1. Cone traps used for covering cowpats to collect
horn and other flies that emerged from the pats.


























































Fig. 4-2. Cages used for testing p
predators on horn fly immatur


redation rates of
es.








81

to the trap) to cover the hole to collect horn flies emerged

from the artificial pat. The mouti of the vial was fitted

with a saran screen funnel to prev nt the flies from going

back to the container. The cages ere maintained at 270C,

50-60% RH and a 14:10 (L:D) photop riod. On the 8th day

after the eggs were seeded, all th containers were covered

by a piece of black polyethylene t rough which holes (5.2

cm) were cut to expose the vials t the light. Attracted by

light, the flies went to the funne led vial soon after they

emerged. Replicates for treatment with one beetle were 10

with P. lonQicornis, 14 with P. fl volimbatus, 10 with P.

ventralis, 7 with P. hepaticus, an 3 for control,

respectively; replicates for treat ents with two beetles

were three each for P. longicornis P. flavolimbatus, and P.

hepaticus.

Differences in mean horn fly mergence between

treatments were analyzed by a one-way completely randomized

analysis of variance (ANOVA), and the significance of

differences between means was test d by Duncan's multiple

range test (SAS 1990). The numbers of adult horn flies

survived of predation were transfo med by log (N+1) to

reduce heteroscedasticity (Dowdy ard Wearden 1983) before

ANOVA was conducted.










Results and Discussion



Laboratory Horn Fly Survival

Percentage hatches of horn fl eggs were from 78 to 94%

(avg. 85.7 4.1%) when the eggs w re distributed on paper

towel strips which were inserted i to the artificial pat for

24 hrs. The percentage survival o the eggs to the adult

stage was from 22 to 41% [29.8 6 15 (SE)].



Fauna-Caused Field Horn Fly Mortalities

1992 July trial

Horn flies started to emerge on the 9th day after their

eggs were seeded under the artificial pat. Most of the

flies emerged on the 10th and 11th day. A few emerged on

the 12th and 13th day. An average of 2.9 0.95 (SE) horn

flies emerged from the pats covere by emergence traps to

exclude other arthropods from arri ing in the dung (range 0-

8); an average of 0.7 0.3 (SE) flies emerged from the pats

exposed to other arthropods (range 0-3). An independent

Student's t-test showed that the m an number of horn flies

that emerged from uncovered pats w s significantly smaller

than that emerged from covered pats (t(18) = 2.51, P < 0.05;

Fig. 4-3). Horn fly numbers were reduced by an average of

75.86% in the uncovered pats compared with the covered ones.










1992 August trial

An average of 3.0 0.56 (SE) horn flies emerged from

the pats covered by emergence traps to exclude other

arthropods from arriving in the dun g (range 0-5); an

average of 1.0 0.26 (SE) flies em erged from the pats

exposed to other arthropods (range 0-2). An independent

Student's t-test showed that the mean number of horn flies

emerged from uncovered pats was significantly smaller than

that emerged from covered pats (ti = 3.25, P < 0.01; Fig.

4-3). Horn fly numbers were reduced by 66.7% in the

uncovered pats compared with those in the covered pats.

Combined data from July and August 1992 showed that the dung

arthropod community caused 71.26% ortality to immature horn

flies in artificially formed cowpais in north-central

Florida.

The mortality of immature horn flies caused by the dung

fauna in the present study was lower than mortalities given

in previous reports. Contributions of the arthropod

community in pastures to horn fly ortality were reported as

87.9% (Roth 1989) and 90% (Kunz et al. 1972) in Texas, 97.7%

in Missouri (Thomas and Morgan 197 b), and 79-84% in

Australia (Fay et al. 1990).

Horn fly survivorship was lower in the field (covered

pats) than in the laboratory. One reason was infestation by

the red imported fire ant (RIFA), which arrived within 24










after the pats were prepared. RIFA workers could also

infest the cone-trap covered pats hrough underground

tunnels. RIFA has a multiple-queen system and its nest

consists of multiple mounds and underground tunnels (Oi et

al. 1994). The fire ants built up small new colonies in the

heavily-infested pats before the horn fly finished its

immature development. Damaged adutt horn flies were

observed in the vial screwed on the top of the cone traps

covering the pats in both treatments and I attribute this

damage to RIFA. The quantitative effect of RIFA on horn fly

populations will be reported in th next chapter. Another

reason for poor survivorship in the field was possibly that

the laboratory colony had been maintained for a long time

leading to genetic changes during the fly's domestication

(Bartlett 1984), and flies were no longer adapted to the

field environment. Desiccation plus high ambient

temperature during the day might be the primary abiotic

factors which decrease immature horn fly survivorship in the

dung.



Predation Rates of Predators on Horn Fly Immatures

Philonthus spp.

Members of the genus Philonthus have been tested as

effective predators of horn flies Thomas and Morgan 1972b;

Harris and Oliver 1979; Roth 1982; Fincher and Summerlin





















July
August







b
R T


Covered










Fig. 4-3. Horn flies emerged fr(
cowpats in a north central I
(n = 10 for each treatment)
indicate significant differ


Uncovered










)m covered and uncovered
Plorida pasture in 1992
Different letters
nces (P < 0.05).


4-



3-



2-


1-



0-


A










1994). During the field survey of the arthropod community

in the present study, specimens of six species of Philonthus

were found in the dung. Living specimens of five species

were colonized in the laboratory for predation rate tests:

P. flavolimbatus, P. hepaticus, P. lonQicornis, P. sericans

and P. ventralis.

Under laboratory conditions. Adults of all five

Philonthus species preyed on horn fly eggs and early instar

larvae. P. longicornis (Fig. 4-4) preferred fly eggs to

larvae; P. ventralis preferred larvae to eggs; the other

three species preyed about equally on eggs and larvae (Fig.

4-5). P. longicornis was observed to consume as many as 6-7

horn fly eggs per minute without interruption; it chewed

the fly egg chorion into shapeless pieces while ingesting

the fluid. Victimized fly larvae had only the

cephaloskeleton and the sclerotize structures of the

posterior spiracle (peritreme, button and spiracular slits)

left on the paper towel or on the cotton ball in the Petri

dish. ANOVA showed that predation rates by these five

Philonthus species on combinations of horn fly eggs and

larvae were significantly different (F (4, 136) = 259.2, P <

0.01; Fig. 4-5). The predation rate of P. longicornis

(120.1 /day) was significantly higher (P < 0.05) than those

of the remaining four species, followed by P. ventralis

(38.9/day). The predation rates of P. flavolimbatus, P.

sericans and P. hepaticus were not significantly different










(P > 0.05). Most Philonthus adults lived 2 to 3 months

under laboratory conditions and they consumed numerous horn

fly immatures during their lives.

Larvae of all five species preyed on horn fly eggs and

larvae. P. lonQicornis had the highest rate, followed by P.

ventralis; P. hepaticus had the lowest rate. P.

longicornis preferred eggs over larvae, but P. ventralis

preferred larvae over eggs. The other three species preyed

on a slightly higher number of fly larvae than of eggs

(Table 4-1). The larvae of all fLve Philonthus species

began to prey on horn fly eggs or Larvae soon after they

hatched. Predation rates then increased, peaking on the 6-

7th day for P. ventralis, and the 4th or 5th day for the

remaining species. P. longicornis preyed on similar numbers

of fly eggs and larvae on the firs day, but on more larvae

than eggs on the following days; P flavolimbatus preyed

primarily on larvae on the first two days, on more eggs

than larvae on the 3-5th day, and then on more larvae than

eggs; P. sericans preyed primaril on larvae for the first

three days after hatching, then on more eggs than larvae

during the following days; and P. hepaticus preyed only on

fly larvae for the first two days, on a few eggs on the 3rd

and 4th day, and then on more eggs than larvae (Fig. 4-6).

All the larvae stopped feeding 1-2 days before they pupated.

When five newly hatched larva of P. longicornis were

confined individually in Petri dis es and provided manure




Full Text
Roth, J.P. 1982. Predation on the h
irritans (L.), by three Philon
Entomol. 7: 26-30.
Roth, J.P. 1983. Compatibility of c
fimicolous staphylinids as bio
the horn fly, Haematobia irrit
Muscidae). Environ. Entomol. 1
Roth, J.P. 1989. Field mortality of
unimproved central Texas pastu
98-102.
Roth, J.P., G.T. Fincher and J.W. S
163
orn fly, Haematobia
thus species. Southwest,
oprophagous scarabs and
logical control agents of
ans (L.) (Diptera:
2: 124-127.
the horn fly on
re. Environ. Entomol,
18:
ummerlin. 1983
Competition and predation as mortality factors of the
horn fly, Haematobia irritans (L.)(Diptera: Muscidae)
in a central Texas pasture habjitat. Environ. Entomol,
12: 106-109.
Bay
Roth, J.P., A. Macgueen and D.E
and predation as mortality fac
Haematobia irritans exigua. in
Southwest. Entomol. 13: 119-12
Roth, J.P., A. Macqueen and D.E. Ba
the introduced phoretic mite,
(Acari: Macrochelidae), on buf
irritans exigua (Diptera: Muse
Environ. Entomol. 17: 603-607.
Roth, J.P., A. Macqueen and D.E. Ba
introduced scarab beetles (Col
coprophilic insects in central
Entomol. 20: 909-914.
Sanders, D.P. and R.C. Dobson. 1966
associated with bovine manure
Soc. Am. 59: 955-959.
Sanders, D.P. and R.C. Dobson. 1969
biology of the horn fly. J. Ec
1366 .
SAS Institute. 1990. User's Guide:
ed. SAS Institute, Cary, NC.
Schmidt, C.D. 1983. Production of h
cattle on three different diet
10: 279-282.
1988a. Scarab activity
tors of the buffalo fly,
central Queensland.
4 .
y. 1988b. Predation by
Macrocheles peregrinus
falo fly, Haematobia
idae), in Australia.
y. 1991. Effect of
eoptera: Scarabaeinae) on
Queensland. Environ.
The insect complex
in Indiana. Ann. Entomol,
Contributions to the
on. Entomol. 62: 1362-
Statistics. Ver. 6, 4th
orn flies in manure from
s. Southwest. Entomol.


Table 3-4 (continued)
Trapped
Extracted
Cicindelidae
Cicindela puntulata Olivier
4
C. scutellaris Dejean
1
Megacephala virginica L.
1
Scolytidae (1 sp.)
27
++
Ptiliidae (1 sp.)
32
++
Elateridae (3 spp.)
13
Anthicidae
Anthicus sp.
6
+
* Carabidae were determined by P. '
Choate
and M. Thomas;
Histeridae by P. Kovarik; Hydrophilidae by A. Smetana; and
Cicindelidae by P. Choate.


150
Dalton, L.W., H.G. Kinzer, J.M. Ree
Host location in the horn fly:
vapor and cow-produced odors i
Entomol. 3: 147-153.
Depner, K.R. 1961. The effect of t
and diapause of the horn fly,
(Diptera: Muscidae). Can. Ente
ves and J.W. Atmar. 1978.
Role of heat, C02, water
n attraction. Southwest.
emperature on development
Siphona irritans L.
mol. 93: 855-859.
Depner, K.R. 1968. Hymenopterous parasites of the horn fly,
Haematobia irritans (Linnaeus), in Alberta. Can.
Entomol. 100: 1057-1060.
Dobson, R.C., F.W. Kunz and D.P. Sa
of horn flies to testosterone
Entomol. 63: 323-324.
nders. 1970. Attraction
treated steers. J. Econ.
Boube, R.C. 1986. biological control
Australia: the potential of th
fauna, pp. 16-34. In R.S. Patt
(eds.): Biological Control of
Soc. Am. Mise. Publ. 61, Colle
of the buffalo fly in
e Southern Africa dung
erson and D.A. Rutz
Muscoid Flies. Entomol.
ge Park, MD.
Doube, B.M. and A. Macqueen. 1991.
dung beetles in Queensland: Th
specificity. Entomophaga 36: 3
Doube, B.M., A. Macqueen and K.A. H
the predatory activity of Macr
(Acarini: Macrochelidae) on tw<
fly (Diptera: Muscidae). pp. 1
and D.A. Rutz (eds.): Biologic
Flies. Entomol. Soc. Am. Mise.
MD.
Establishment of exotic
e role of habitat
35-360.
uxham. 1986. Aspects of
cheles perearinus
o species of Haematobia
32-141. In R.S. Patterson
al Control of Muscoid
Publ. 61, College Park,
Doube, B.M. and F. Moola. 1988. The effect of the activity
of the African dung beetle Catharsius tricornutus
DeGeer (Coleptera: Scarabaeidae) on the survival and
size of the African buffalo fly, Haematobia thirouxi
potans (Bezzi) (Diptera: Muscidae), in bovine dung in
laboratory. Bull. Entomol. Re
s. 78: 63-73.
Dowdy, S.M. and S. Wearden.
Wiley, New York.
1983. Statistics for research.
Drea, J.J. 1966. Studies on Aleocha
Staphylinidae), a natural enem
Econ. Entomol. 59: 1368-1373
ra tristis (Coleptera:
y of the face fly. J.


No. Beetles Collected / Trap (Avg. +/- SE)
43
Fig. 3-2. Seasonal distribution of five families of
Coleptera most commonly collected by trapping (Mean
Se). N = 10 X 2 for May to October and N = 10 for the
remaining months.


control (F(1 22) = 4.86, P
128
the trial of 1992. The mean number of staphylinids
collected in the Amdro-treated are^ was significantly
greater than that collected in the
< 0.05). More hydrophilids were collected in the Amdro-
treated area than in the control area (F(1 22) = 2.01, P <
0.05) .
Among the staphylinids collected in pitfall traps, the
mean number of Philonthus spp. collected in the Amdro-
treated area (13.3 7.4) was significantly greater (t(22) =
2.22, P < 0.05) than that in the control area (3.5 7.2).
Numbers of all four Philonthus spec
area were greater than those in the
8). Other staphylinids including 0
ies in the Amdro-treated
control area (Table 5-
xvtelus incisus.
Neohypnus pusillus. and Tinotus spp
the Amdro-treated area. These speci
predators of horn fly immatures (ch
Among Diptera, mean numbers of
collected in the Amdro-treated area
greater than those in the control a
numbers of the other families of Di
Amdro-treated area were not signifi
control area (P > 0.05).
The main species of arthropods
. were more abundant in
es were tested as
apter 4).
muscids and sepsids
were significantly
rea (P < 0.05). Mean
ptera collected in the
:antly different from the
extracted from cowpats
in both 1992 and 1993 included Cercyon spp. (Hydrophilidae),
Tachvs sp. (Carabidae) Brontaea debilis and B_;_ cilfera
(Muscidae), Ravinia derelicta. R, f
loridensis and Helicobia


162
Paulian, R. 1941. Les premiers tat
s des Staphylinoidea.
tude de morphologie compare.
Mm. Mus. Natn. Hist.
Nat. Paris (n. ser.) 15: 1-361

Peck, 0. 1974. Chalcidoid (Hymenopt
era) parasites of the
horn flv, Haematobia irritans
(Diptera: Muscidae), in
Alberta and elsewhere in Caad
a. Can. Entomol. 106:
473-477.
Peschke, K. and D. Fuldner. 1977. U
ebersicht und neue
Untersuchungen zur Lebensweise
der Parasitoiden
Aleocharinae (Coleptera: Stap
hylinidae). Zool. Jb.
Syst. 104: 241-262.
Pickens, L.G. 1981. The life histor
y and predatory
efficiency of Ravinia Ihermini
eri (Diptera:
Sarcophagidae) on face fly (Di
ptera: Muscidae). Can.
Entomol. 113: 523-526.
Pimentel, D. 1955. Relationship of
ants to fly control in
Puerto Rico. J. Econ. Entomol.
48: 28-30.
Poorbaugh, J.H., J.R. Anderson and
J.F. Burger. 1968. The
insect inhabitants of undistur
bed cattle droppings in
northern California. Calif. Ve
ctor Views 15: 17-36.
Porter, S.D. and D.A. Savignano. 19
90. Invasion of polygyne
fire ants decimates native ant
s and disrupts arthropod
community. Ecology 71: 2095-21
06 .
Porter, S.D. and W.R. Tschinkel. 19
87. Foraging in
Solenopsis invicta (Humenopter
a: Formicidae): effects
of weather and season. Environ
. Entomol. 16: 802-808.
Quisenberry, S.S. and D.R. Strohbeh
n. 1984. Horn fly
(Diptera: Muscidae) control on
beef cows with
permethrin-impregnated ear tag
s and effect on
subsequent calf weight gains.
J. Econ. Entomol. 77:
422-424.
Reagan, T.E., G. Coburn and S.D. He
nsley. 1972. Effect of
mirex on the arthropod fauna o
f a Louisiana sugarcane
field. Environ. Entomol. 1: 58
8-591.
Riley, C.V. 1889. The horn fly. Ins
ect Life 2(6): 93-103.
Roberts, R.H. and W. A. Pund. 1974.
Control of biting flies
on beef steers: Effect on per
formance in pasture and
feedlot. J. Econ. Entomol. 67:
232-234.


138
There are over 400 species of
arthropod species
reported associated with cattle dur
g in the USA (Blume 1985;
Harris and Blume 1986; Fincher 1990
). However, the dung
arthropod fauna in this country is
impoverished compared
with that in Europe where the horn
fly originated but causes
no serious damage to the cattle inc
Lustry. Staphylinidae,
for example, the most important pre
.dators of the horn fly
(Hammer 1941; Blume et al. 1970; Me
icqueen and Beirne 1975;
Thomas and Morgan 1972b), have 43 £
.pecies reported in the
USA (Fincher 1990), while 133 speci
.es were reported from
cattle dung in a study in Finland
Loskela 1972). There are
over 5,000 species of scarab beetle
is worldwide (Skidmore
1991; Hanski 1991), but only over !
.00 species are recorded
in the USA (Blume 1995; Harris and
Blume 1986; Fincher
1990).
The most diverse and abundant
order was Coleptera, in
which Staphylinidae were ranked the
i first in species
richness (44 species), followed by
Scarabaeidae (27 species)
and Carabidae (20 species). In ni
imbers of specimens
collected, Staphylinidae were the n
lost abundant, followed by
Scarabaeidae, Hydrophilidae, Histei
~idae, and Carabidae.
Members of Staphylinidae, Hydrophi]
Lidae and Histeridae have
been documented as predators of imi
nature horn flies (Hammer
1941; Blume et al. 1970; Macqueen
ind Beirne 1975; Thomas
and Morgan 1972b), and Scarabaeidae
; as competitors (Anderson
and Loomis 1978; Blume et al. 1973,
Bornemissza 1970, 1976;


60
months of the year. A^_ viridis were collected in the
summer, and Th. intersectus and P^_ sublaevis were collected
during the summer and fall.
Most species in this family were collected during the
day time.
Hydrophilidae. Sixteen species of Hydrophilidae have
been reported associated with cow dung in the continental
USA (Fincher 1990), but only four species were collected in
the present survey. Sphaeridium scarabaeoides (L.) has been
considered to be an important predator of horn flies (Bourne
and Hays 1968; Hammer 1941; Macqueen and Beirne 1975; Mohr
1943; Poorbaugh 1966; Sanders and Dobson 1966). S_;_ lunatum
were commonly collected. It is similar in size to S.
scarabaeoides and these two species may have similar
predation potential against the horn fly. Cercvon
variecratus and Ch. atricapillus were abundant (Table 3-4)
but they have not been considered to be predators (Sanders
and Dobson 1966; Hafez 1939; Merritt 1976; Thomas and Morgan
1972b).
More Hydrophilidae were col
winter than in the summer and fall
Histeridae. Twenty-two sped
reported from dung in the continen
Only four species of these beetles
lected in the spring and
(Fig. 3-2).
es of Histeridae have been
tal USA (Fincher 1990).
were collected in this
survey. The most abundant was Hister coenosus, which is
medium-sized and has been considered to be the most


51
tetracariatus Block and Atheta sp. as small predators but
did not present evidence. It is reported that Platystethus
spiculus (Palomino and Dale 1989), Ft_ americanus (Mohr 1943;
Cervenka and Moon 1991) and Falaqria dissecta (Valiela 1974)
are predators of flies. These species, however, were not
abundant in this study.
Staphylinidae were collected year round. Numbers were
low during the winter, built up slowly in the spring, and
peaked in the late summer or fall. The seasonal abundance
for 1993 showed bimodal peaks in July-August and November
(Fig. 3-2). The first peak was mainly composed of Oxvtelus
incisus. and Aleochara notula (Fig. 3-4); the second peak
was mainly composed of Anotvlus spp., Tinotus spp. and
Acrotona hebeticornis (Fig. 3-4).
The number of species of Staphylinidae was low in the
spring, increased in the early summer, peaked in June-August
(14-16 species), and then declined (Fig. 3-3). Different
staphylinid species were collected during the different
seasons of a year. Anotvlus insicmitus and A_;_ nanus were
mainly collected during the winter and spring. Meronera
venustula and Mycetoporus flavicollis were mainly collected
in the spring. Tinotus brunnipes, T. amplus and Acrotona
hebeticornis were mainly collected during the winter.
Atheta sp. were collected each month. Philonthus spp., A.
notula, and CA. incisus were mainly collected during the
summer and fall. Most of the remaining species were


145
of positive effect of fly populations by providing oxygen
supplies through the tunnels (Valiela 1974). Hymenopterous
parasitoids seem to attack few fly puparia in pastures, so
their use as biological control agents in this habitat may
be impractical; this is not to say that they may not be
effective where cattle are crowded, such as in stockyards.
Importation and release of additional effective
habitat-specific or host-specific predators into the
existing fauna should be attempted. No attempt has yet been
made to do this in Florida. The higher levels of mortality
inflicted on horn fly by predators in some other parts of
the USA suggest that some component of the predatory fauna
may be lacking in Florida. Insects that prey only on
immature horn flies are yet unknown and may not exist. But
the major habitat of some of the predators encountered in
this study, and of others known from elsewhere, is bovine
dung. One method of selection would be to find predators
that might complement these detected in this study, for
example by having their peak activity at some time of year
that is not well "covered" by existing predators. Another
avenue would be to find species of Aleochara that would
attack horn fly pupae more readily than does Aleochara
notula.


79
Fig. 4-1. Cone traps used for covering cowpats to collect
horn and other flies that emerged from the pats.
vv


105
eggs and early instar larvae soon after hatching.
Consumption rate increased, reaching the highest level on
the 11th and 12th day, and then declined (Fig. 4-9). The
larva preferred fly larvae over eggs, and preyed only on fly
larvae before the prepupal stage (f'ig. 4-9). Larvae
consumed a mean 9.0 fly eggs and 8^.5 larvae during their
active stage (Table 4-2). All the
were shown to prey on both horn fl^ eggs and larvae. They
had higher predation rates than did Oxytelinae and
Paederinae, but lower than Philonthus species. N_;_ pusillus
species of Xantholininae
may be the most important predator
subfamily, because it was more abun
species.
Tachvporinae
Mycetoporus flavicollis. Adul
observed to prey on fly larvae only
consumed an average of 2.89 2.51
day (range 0-9).
of horn flies in this
dant than the other
ts of this species were
Each beetle (n = 9)
(SD) horn fly larvae per
were tested for
Aleocharinae
Four species in this subfamily
predation. Falacrria dissecta was not observed to prey on
horn fly eggs and larvae. Atheta sp. and Tinotus amplus
preyed on horn fly larvae only. Each Atheta sp. adult (N =
38) consumed an average of 2.92 1.42 (SD) larvae (range 1-


61
efficient histerid in reducing horn fly populations
(Summerlin et al. 1982, 1984c, 199,0) The other three
species were less common, of which Saprinus pennsvlvanicus
and Phelister haemorrhous have also been reported to be
predators of horn fly immatures (Summerlin et al. 1982,
1991).
Diptera
In total, 35 species of Diptera belonging to 19
families were collected (Table 3-5). Abundant species of
higher dipterans included Haematobia irritans. Brontaea
debilis, B_j_ cilfera and Neomvia cornicina in the family
Muscidae; and Ravinia derelicta. r[ floridensis. Helicobia
morionella and Oxvsarcodexia ventricosa in the family
Sarcophagidae. Abundant lower dung-inhabiting dipterans
included Coproica sp. (Sphaeroceridae), Palaeosepsis
insularis (Sepsidae), Aphodiplosis triangularis and
Lestodiplosis sp. (Cecidomyiidae), Bradysia coprophili
(Sciaridae) and Svlvicola notialis (Anisopodidae).
H. irritans arrives earliest at the fresh dung and
emerges earliest of all the flies from the dung. Previous
study shows that this fly was active all year round in north
central Florida (Wilkerson 1974). The fly numbers were
observed to be low during the early spring, became high in
May and peaked in August to September on the cattle in 1993.
The adults were extracted from dung pats collected in


1977). However, Howard and Oliver
Scarabaeidae (Aphodininae) were aff
poisoned insects in pastures treate
In the 1992 experiment populations
significantly lower in the Amdro-tr
those in the fire ant infested area
has a poisonous effect on scarab be
Howard and Oliver (1978)
Previous studies showed that R
on staphylinid beetles (Howard 1975
1978), which was confirmed by the p
present study showed that S^. invict
Staphylinidae, especially Philonthu
showed negative effects on hydrophi
even though these beetles were not
treated and the control areas. Thi
impede the establishment of insects
biological control of horn and othe
However, developing fly larvae in t
vulnerable to these ants. S. invict
136
(1978) showed that
ected by scavenging dead
d with Mirex ant bait,
of Scarabaeidae were
eated area than were
, suggesting that Amdro
etles. This agreed with
IFA had negative effects
; Howard and Oliver
resent study. The
a decreased abundance
s spp. The results also
lid and carabid beetles,
abundant in the Amdro-
s indicates that RIFA may
introduced for
r flies in the dung,
he dung-pats are highly
a forage night and day,
and are active throughout the fly s
1984a). In general, well-establish
imported fire ant should help suppr
irritans and the other muscid flies
sarcophagid flies Ravinia spp. and
the ants reduce the numbers of dung
suppress populations of these flies
eason (Summerlin et al.
ed populations of the red
ess populations of H.
(Brontaea spp.), and
H. morionella), though
-inhabiting beetles that
through predation.


160
Macqueen, A. and B.P. Beirne. 1975.
insects on production of horn
(Diptera: Muscidae), from catt
British Columbia. Can. Entomol
Influence of other
fly, Haematobia irritans
le dung in south central
. 107: 1255-1264.
Madsen, M., B. Overgaard Nielsen, P
J. Brochner Jespersen, K.M. Va
J.Gronvold. 1990. Treating cat
Effects on fauna and decomposi
Appl. Ecol. 27: 1-15.
. Holter, O.C. Pedersen,
gn Jensen, P. Nansen and
tie with ivermectin:
tion of dung pats. J.
Mank, H.G. 1923. The biology of the
Entomol. Soc. Am. 16: 220-237.
Staphylinidae.
Ann.
Marley,S.E., J.A. Lockwood, R.L. By
1991. Temporal, climatic, and
of dispersal in the horn fly,
(Diptera: Muscidae). Environ.
ford and D.G. Luther,
physiological medication
Haematobia irritans (L.).
Entomol. 20: 1612-1618.
Martin, M.M. and J.S. Martin. 1978.
the midgut of the fungus-growi
natalensis: the role of acquir
Science 199: 1453-1455.
Cellulose digestion in
ng termite Macrotermes
ed digestive enzymes.
McLintock, J. and K.R. Depner. 195
history and habits of the horn
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4
. A review of the life-
fly, Siohona irritans
Entomol. 86: 20-33.
Melvin, R. 1934. Incubation period of eggs of certain
muscoid flies at different constant temperatures. Ann.
Entomol. Soc. Am. 27: 406-410.
Melvin, R. and D.W. Beck. 1931. Len
stages of the horn-fly, Haemat
constant temperature. J. Econ.
gth of the developmental
obia irritans (Lin.) at
Entomol. 24: 330-331.
Merritt, R.W. 1976. A review of the
insect fauna inhabiting cattle
central California. Pan-Pacifi
food habits of the
droppings in north
p Entomol. 52: 13-22.
Merritt, R.W. and J.R. Anderson. 19
different pasture and rangelan
annual dynamics of insects in
Vector Views 24: 43-46.
77. The effect of
1 ecosystems and the
cattle droppings. Calif.
Miller, J.A., D.B. Thomas, A.J. Sie
1984. Estimating numbers of ho
dung. Southwest. Entomol. 9: 2
benaler, and J.J. Matter,
rn fly eggs in bovine
75-280.
Mohr, C.O. 1943. Cattle droppings a
Monogr. 13: 275-298.
s ecological units. Ecol.


22
Fincher (1990) reported 43 species associated with cattle
dung in the continental USA. The important subfamilies
include Aleocharinae, Oxytelinae, Paederinae, Staphylininae
and Tachyporinae (Hunter et al. 1991). Staphylinid beetles
prey on the dung fauna as adults and as larvae (Drea 1966;
Fincher and Summerlin 1994). Predation by Staphylinidae on
horn fly eggs and early instar larvae has been reported for
Philonthus (Harris and Oliver 1979; Hunter et al. 1989;
Macqueen and Beirne 1975; Roth 1982; Thomas and Morgan
1972b), and Aleochara (Klimaszewski and Blume 1986; Harris
and Blume 1986) .
Members of the genus Philonthus are important predators
of dung-inhabiting flies as adults and larvae (Hammer 1941;
Laurence 1954; Sanders and Dobson 1966; Valiela 1969a;
Macqueen and Beirne 1975; Wingo et al. 1974; Wharton 1979).
Fifteen species of Philonthus were reported to inhabit
cattle dung in this country (Fincher 1990). P. cruentatus
(Gmelin) was shown to be one of the most effective predators
on immature stages of the horn fly (Thomas and Morgan
1972b). P. flavolimbatus Erichson was shown to reduce horn
fly production by 69% and 86 when 2 and 4 beetles were
tested for each trial (Roth 1982). Horn flies were reduced
by 91% and 99% when 5 and 10 beetles of this species were
tested for each trial (Harris and Oliver 1979). P.
flavolimbatus and P. cruentatus were found to prey
primarily on the egg stage, while P. rectanqulus Sharp is a


Bennett, P.
Choate, G.
A.
Dahlem, D.
Deonier, R. J. Gagn,
w.
L. Grogan
, V. Gupta,
D.
Habeck, M.
E. Hennessey, T. J.
Henry, P. W.
Kovarik, G
. W
. Kranz, J.
McNamara, F. Mead, A.
S.
Menke, K.
Nguyen, M.
E.
Schauff, P
. Skelley, A. Smetana,
R.
J. Snider
, G. Steck,
G.
B. Edwards
, J. Watts, M. Thomas,
D.
Williams,
W. Wirth.
I am appreciative of members of the insect biocontrol
lab, including J. Cicero, R. Coler, R. Hemenway, P. Parkman
and S. Wineriter for helping me and creating a comfortable
and friendly working atmosphere; D. Simon, H. Brown and J.
Okine in Dr. Butler's lab for helping me to collect immature
horn flies and offering me fly chow essential for the
experiments on predation.
Special thanks go to my family. During the four years'
study my wife, Yanfen Chen, endured and sacrificed much
(especially when she was pregnant with our son) to ensure
time for my study. I appreciate my daughter, Wenli, who
exercised a great deal of tolerance and understanding of my
absence.


68
Table 3-7. Miscellaneous invertebrates collected in pastures
in north central Florida. The occurrence codes for
species not enumerated: (+) rare, (++) common, and
(+++) abundant.
Trapped Extracted
Othoptera
Gryllidae
Gryllus rubens Scudder
++
+
G. firmus Scudder
Gryllotalpidae
++
+
Scapteriscus vicinus Scudder
+
+
Homoptera
Cercopidae
28
Prosapia bicincta (Say)
Cicadellidae
+
+
Exitianus exitiosus (Uhler)
+
+
Planicephalus flavicosta (Stl)
Delphacidae
+
+
Toya andromeda (Van Duzee)
+
+
T. propinqua (Fieber)
+
Hemiptera
162
31
Nabidae (1 species)
Cydnidae
+
Pangaeus bilineatus (Say)
Reduviidae
++
+
Xylocoris galactinus Reuter
Anthocoridae
++
+
Lasiochilus pallidulus Reuter
++
+
Orius sp.
Lygaeidae
+
Pachybrachius basalis (Dallas)
+
Paromius longulus (Dullas)
+
+
Geocoris uliginesis (Say)
+
+
Blissus insularis Barber
+
+
Lepidoptera
Tineidae (1 species)
Pyralidae
1
Herpetogramma
phaeopteralis (Guene)
Noctuidae
8
Spodoptera frugiperda (Smith)
42
++


102
Table 4-2. Predation by N. pusillus larvae and adults on
eggs and larvae of horn flies
conditions.
under laboratory
Predator
stages
Prey
stages
No.
predators
observed
No. of prey consumed
Range Mean SD
Larva(whole
active stage)
Egg
19
0-25
9.0
8.6
Larva
19
40-140
83.5
21.5
Adult (each
day)
Egg
29
0-13
4.2
3.1
Larva
29
0-17
5.9
5.1




56
small beetles swarmed into the dung pat within a few house
after the dung was deposited. One day later the dung pat
was mostly consumed and totally broken up, and then became
dry in 4-5 days. Flies and other immature insects were not
found abundant during this time. 0^_ qazella. which were
released in Texas in 1972 and found in Florida in 1975
(Fincher 1990), were abundant onlyl in July and August. It
was reported that this beetle removed over 80% of dung from
the soil surface during its peak activity in Texas (Harris
1981). A similar situation was observed but not measured in
July and August in the present survey. 0_^ taurus. too, is
not a native species (Fincher 1990), but its numbers were
much lower than those of Cb_ qazella and the other native
Onthophagus species.
The remaining five species of Onthophagus, which were
abundant during the fall and early winter (Fig. 3-5), were
less important than A_^ lividus and 0^_ qazella in
decomposition of dung.
No scarab beetles were collected in January, but the
number of species increased from February, was highest in
June to August (14-15 species), then declined. There was a
second peak in November (Fig. 3-3)1.
Most 0_;_ qazella. other Onthophagus spp. and A^_ lividus
were collected during the daytime.
Though small numbers of Ataenlus spp. and Aphodius spp.
were extracted 1-3 weeks after the pats were collected and


No. Horn Flies Emerged (Mean +/- Se) / Pat
85
Covered
Uncovered
Fig. 4-3. Horn flies emerged from covered and uncovered
cowpats in a north central Florida pasture in 1992
(n = 10 for each treatment). Different letters
indicate significant differences (P < 0.05).


Figure Page
4-8. Predation of P. lonqicornis on horn flies at
different prey densities under simulated field
conditions 95
4-9. Daily predation rates of N. pusillus larvae on horn
fly eggs and larvae under laboratory conditions 103
4-10. Aleochara notula preying on horn fly eggs under
laboratory conditions 104
5-1. Fire ants preying on horn fly larvae
in the laboratory 118
x


151
Drummond, R.O., G. Lambert, H.G. Sir
Terill. 1981. Estimated losses
In D. Pimentel (ed.) CRC Handb
agriculture, vol.l, pp. 111-12
Fla.
alley, Jr. and C.E.
of livestock to pests,
ook of pest management in
7. CRC Press, Boca Raton,
Drummond, R.O., J.E. George and S.E. Kunz. 1988. Control of
arthropod pests of livestock: A review of technology.
CRC Press, Boca Raton, Fla.
Eddy, G.W., A.R. Roth and F.W. Plapp Jr. 1962. Studies on
the flight habits of some marked insects. J. Econ.
Entomol. 55: 603-607.
Endris, R.G., D.G. Young and P.V. P
Ultrastructural comparison of
five Lutzomvia species (Dipter
Entomol. 24: 412-415.
erkins. 1987.
egg surface morphology of
a: Psychodidae). J. Med.
Escher, R.L. 1977. Pupal parasites
Haematobia irritans (L.) (Dipt
central Florida. Thesis, Dept.
Florida, Gainesville, 88 pp.
of the horn fly
era: Muscidae) in north-
Entomol & Nematol. Univ.
Eschle, J.L., J.A. Miller and C.S.
growth regulators and sterile
horn flies. Nature 265: 325-32
Schmidt. 1977. Insect
males for suppression of
6 .
Esser, R. 1980. Nematomorpha. Nema
Dept. Agrie. Cons. Serv. Gaine
Fay, H.A.C. 1986. Fauna-induced mor
thirouxi Potans (Bezzi) (Dipte
dung in relation to soil and v
149. In R.S. Patterson and D.A
Control of Muscoid Flies.
Publ. 61.
tology circular. 70. Fla.
sville.
tality in Haematobia
ra: Muscidae) in buffalo
egetation type. pp. 142-
Rutz (eds): Biological
Entomol. Soc. Am. Mise.
Fay, H.A.C. and B.M. Doube. 1983. T
coprophagous and predatory bee
immature stages of the African
thirouxi potans. in bovine dun
460-466.
he effect of some
ties on the survival of
buffalo fly, Haematobia
g. Z. Ang. Entomol. 95:
Fay,
H.A.C., A. Macqueen and B.M. D
fauna on mortality and size of
(Diptera: Muscidae) in natural
and South Africa. Bull. Entom
oube. 1990. Impact of
Haematobia spp.
dung pads in Australia
ol. Res. 80: 385-392.
Ferrar, P. 1975.
beetles. J.
Preamble for symposium on effects of dung
Appl. Ecol. 12: 819-821.


Table 3-5. Dptera collected in pastures in north central
Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and (+++) abundant.
Trapped
Extracted
Muscidae
Haematobia irritans L.
0
226
Brontaea debilis (Williston)
++
375
B. cilfera (Malloch)
++
198
Pseliphephila sp
+
+
Neomyia cornicina (F.)
++
497
Sphaeroceridae
Coproica sp.
+ +
2221
Sepsidae
Palaeosepsis insularis (Williston
) +++
1358
Ephydridae
Discocerina obscurella (Fallen)
2
Sarcophagidae
Ravinia derelicta (Walker)
++
226
R. floridensis (Aldrich)
++
79
Udamopyga niagrana (Parker)
2
+
Oxysarcodexia ventricosa (Wulp)
++
38
Helicobia morionella (Aldrich)
++
33
Cecidomyiidae
Aphodiplosis triangularis (Felt)
++
1142
Lestodiplosis sp. (new)
+++
760
Neolasioptera sp.
++
620
Aprionus sp.
++
18
Bibionidae
Plecia nearctica Hardy
+
+
Dolichopodidae
Sciapces sp.
++
+
Syntormor sp.
35
Sp. 3
++
+
Empididae
Drapetis vanthopoda Will.
++
47
Milichiidae (1 species)
4
++


96
more effective against horn fly eggs than larval and pupal
stages, which agrees with the result from the present study.
Some Philonthus species, such as P, lonoicornis
(present study) and P. flavolimbatus (present study and
Harris and Oliver [1979]) seem to have the potential to
destroy all horn fly eggs and larvae. That they do not do so
in the field suggests that their population densities or
dispersion are inadequate, or that
alternate prey distracts them from
horn fly immatures. Nevertheless,
important mortality on horn fly.
their feeding or
complete elimination of
they may inflict very
Oxvtelinae
Platvstethus americanus. Ten
were confined individually in Petri
with horn fly eggs and first instar
manure. Each beetle consumed 1.5
larvae of the horn fly per day (ran
the pharyngeal sclerites of the lar
beetle eggs were found in the dishe
adults were offered cow manure for
chambers, they survived and the fern
laid eggs in them. This suggests t
prey on horn fly larvae for nutrien
chambers for oviposition.
adult P. americanus
dishes and provided
larvae for food, without
+ 1.02 (SD) first instar
ge 0-4/day), leaving only
vae in the dish; but no
s. However, when these
food in the rearing
ales made chambers and
hat adult P. americanus
ts and need dung to form
When five larvae of P. america:
nus were confined in


4


71
by insect colonizers, including scarab and staphylinid
beetles, and muscid, sarcophagid, sphaerocerid and sepsid
flies. These mites were frequently found on the dung surface
in the field. Four families of mites, Anistidae (one
species), Parasitidae, Macrochelidae, and Oribatidae, were
collected, among which members of Parasitidae and
Macrochelidae have been reported to be predators of fly
immatures (Valiela 1974; Krantz 1983; Halliday and Holm
1987; Anderson 1983). Macrochelids have been considered to
have biological control potential of house, bush, face and
horn flies (Axtell 1963; Krantz 1983; Halliday and Holm
1987; Anderson 1983), in particular, M^ muscaedomesticae
(Axtell 1963) and it peniculatus (Roth et al. 1988b;
Halliday and Holm 1987). Oribatids occur in the old dung and
are thought to feed on the dung or on associated fungi
(Skidmore 1991).
Araneae. Spiders were often found in dung. They are
general predators, preying on whatever they meet and are
able to handle.
Millipedes. Millipedes are associated with decaying
plant materials, upon which they feed. They were commonly
collected by trapping and extracting and were abundant in
the fall and winter. There are no data on the relationship
between millipedes and flies.
Collembola. Springtails were abundantly collected by
trapping and extracting. They feed on decaying plant


132
Table 5-7. continued.
Arthropod taxa
No. collected
Amdro
Control
Sciaridae
Mean
4.25
8.92
SD
12.01
29.27
Range
0-44
0-106
F 0,
P
>
0.10
Anisopodidae
Mean
1.08
1.17
SD
1.04
2.54
Range
0-3
0-9
F = 0.38,
P
>
0.10
Ceratopogonidae
Mean
2.42
8.33
SD
2.43
18.02
Range
0-7
0-65
F = 0.57,
P
>
0.10
Psychodidae
Mean
1.58
2.5
SD
2.29
2.6
Range
b-8
0-7
F = 2.48,
P
>
0.10
Hymenoptera
(except RIFA)
Mean
2.42
1.50
SD
k .41
3.55
Range
0-16
0-13
F = 0.18,
P
>
0.10


169
Woodruff, R.E. 1973. The scarab bee
Arthropods of Florida and Neig
8. Fla. Dept. Agrie., Divn. PI
Gainesville.
Wu,
W.J. and W.Z. Zhang. 1990. Stud
function of Creophilus maxillo
Staphylinidae) for fly larvae.
Zorka, T.J. and D.E. Bay. 1980. The courtship behavior of
the horn fly. Southwest. Entomol. 5: 196-200.
ties of Florida. Pt. 1.
hboring Land Areas. Vol.
ant Industry,
y on the biology and prey
sus (Coleptera:
Pest Control 6: 19-21.
Zumpt, F. 1973. The stomoxyine bit
(Diptera: Muscidae) Taxonomy,
importance and control measure
Stuttgart, 175 pp.
ing flies of the world,
biology, economic
s, Gustav Fischer Verlag,


Under laboratory conditions t
9
lorn fly larvae can develop
in cattle, bison, sheep, and horse
i manure (Greer and Butler
1973), but in the field larval development takes place only
in cattle manure (Depner 1961, Bruce 1964, Foil et al.
1990). Experiments show that cattle fed a large amount of
grain produce manure which is unusually acid, reducing horn
fly larval survival under laboratory conditions (Schmidt
1983).
Pupa. The pupa is coarctate
and the puparium is dark
brown. It is approximately 3.3 mm
long and 1.4 mm wide. This
stage lasts about 5.5 days under r
atural field conditions.
It can differ by three to four days depending whether the
manure pat is in sunlight or shads
2 (Kunz et al. 1970).
Emergence of the flies from manure
2 occurs from mid to late
afternoon in the summer, but at any time of day in the
spring and fall (Lancaster and Me;
Lsch 1986). Males need one
day less to develop at this stage
than do females (McLintock
and Depner 1954; Hoelscher and Comb 1971). Sex ratio was
reported as 1:1 or 1:1.35 (Glaser
1924; Mohr 1943).
Adult. The adult horn fly is
3 about 4 mm long (Foil et
al. 1990), or about one-third to c
3ne-half the size of the
common house flv. Musca domestica
(L.) (Okine 1991).
Longevity of the horn fly is estimated from 28 days
(McLintock and Depner 1954) to 6-
3 weeks (Bruce 1964). Male
survival rates were reported to bs
2 about 95% of the female
survival rate (Krasfur and Ernst :
L983). Both sexes have


9
6
3
O
8
6
4
2
O
4
3
2
1
O
47
M J J A
Month 1993
4. Seasonal distribution of staphylinid species in
sture A, collected by trapping. N = 10 X 2 for May
October, and N = 10 for the remaining months.


98
average of 1.92 0.98 (SD) horn fly larvae per day (range
1-4). The females oviposited when they were confined
individually in Petri dishes and provided only with horn fly
eggs and larvae for food, without manure. The result showed
that predation on fly larvae is the normal feeding, or at
least adequate behavior of this beetle, and the nutrients
from horn fly larvae are enough for it to survive well and
produce offspring.
Larvae of A_¡_ insignitus (n = 12) were observed to
consume an average of 1.78 1.03 (SD) horn fly larvae per
day, but no eggs. They started to prey on 1st instar horn
fly larvae soon after hatching. Ech larva of A_;_ insignitus
consumed an average of 18.7 8.53
the active stage and then
its nine days of activity. Actually A^_ insignitus killed
more fly larvae than they consumed, especially during the
first two days after they hatch. When only fly eggs and
larvae were offered for food, without manure, the beetle
larvae wandered for two days after
died without pupation. When the larvae were reared with
manure and horn fly eggs and larvae, pupation occurred. The
adults of A_j_ insignitus did not need manure to oviposit. The
larvae, however, needed manure in which to form pupation
cells.
Oxvtelus incisus. Adult O^. iricisus were observed to
(SD) fly larvae during
prey primarily on horn fly larvae.
confined individually in Petri dishes, only two fly eggs
When seven adults were


118
Fig. 5-1. Fire ants preying on hor
laboratory.
n fly larvae in the


and thawed. Selected epigeal arthropods were sampled by-
fresh cattle manure,
ication of Amdro, ten
nure were set up in the
ociated with cattle dung
1) and emergence boxes
in the Amdro-treated and
cone traps on the 8th day
means of pitfall traps baited with
Before and 4-6 weeks after the appl
pitfall traps baited with bovine ms.
grid pattern in the Amdro-treated and control areas. Traps
remained in the pasture for 24 hours, the arthropods were
collected into 70% alcohol, and brought into the laboratory
for identification. Arthropods ass
were sampled by cone traps (Fig. 4-
(Fig. 3-1). In 1992, 12 pats each
the control area were covered with
of the fly egg-seeding for collecting insects flying into
the tube on the top of the trap (chapter 4); 6 pats each in
RIFA-controlled and infested areas
laboratory with emergence boxes to
and other insects. In 1993, few arthropods emerged from the
first set of 12 pats in each treatment area, because of a
continuous heavy rain that drowned
insects. For the second set of 12
covered by corn traps and the other six were brought into
the lab for extraction of arthropods. The numbers of
arthropods from the two sets of pats in each area were
combined for analyses.
Artificial pats were prepared and seeded with horn fly
eggs (100 eggs per pat) by following the method of Thomas
114
were brought into the
extract flies, beetles
most of the immature
pats, six of them were
and Morgan (1972b). Horn fly eggs
were obtained from a


TABLE OF CONTENTS
ACKNOWLEDGMENTS ii
LIST OF TABLES V
LIST OF FIGURES ix
ABSTRACT xi
CHAPTERS
1. INTRODUCTION 1
2. LITERATURE REVIEW 4
History and Importance of the Horn Fly .... 4
History 4
Economic Importance 4
Biology of the Horn Fly 7
Life History 7
Mating and Oviposition 10
Host Orientation and Location 11
Diapause and Dispersal 13
Control of the Horn Fly 14
Chemical Control 14
Genetic, Physical and Immunological
Control 15
Biological Control 16
Dung Arthropod Community 17
Dung Arthropod Community Composition ... 17
Succession of the Community 19
Effect of the Whole Insect Fauna on
Horn Fly Production 20
Predators 21
Staphylinidae 21
Hydrophilidae 23
Histeridae 23
Red imported fire ant 24
Mites 26
Flies 27
Competitors 28
Coprophagous beetles 28
Coprophagous flies 33
iv


155
Harris, R.L. and J.W. Summerlin. 1984. Parasites of horn fly
pupae in east central Texas. Southwest. Entomol. 9:
169-173.
Harvey, T.L. and J.R. Brethour. 1970. Horn fly control with
dichlovos-impregnated strip. J. Econ. Entomol. 63:
1688-1689.
Harvey, T.L. and J.R. Brethour. 1979. Effect of horn flies
on weight gains of beef cattle. J. Econ. Entomol. 72:
516-518.
Harvey, T.L. and J.L. Launchbaugh. 1982. Effect of horn
flies on behavior of cattle. J. Econ. Entomol. 75:25-
27.
Haufe, W.O. 1982. Growth of range c
flies (Haematobia irritans) by
with fenvalerate. Can. J. Anim
attle protected from horn
ear tags impregnated
. Sci. 62: 467-573.
Hays, S.B. and K.L. Hays. 1959. Food habits of Solenopsis
saevissima richteri Forel. J. Econ. Entomol. 52: 455-
457.
Hinton, H.E. 1944. Some general rem
beetles, with notes on the bio
Platvstethus arenarius (Fourcr
Entomol. Soc. London (A) 19: 1
arks on sub-social
logy of the staphylinid
oy). Proc. Royal
15-128.
Hinton, H.E. 1981a.
Press, Oxford.
Biology of Inse
t
Eggs.
Vol 1.
Pergamon
Hinton, H.E. 1981b.
Press, Oxford.
Biology of Inse
ct Eggs. Vol 2. Pergamon
Hoelscher, C.E. and R.L. Combs, Jr. 1971. The horn fly. III.
Sex ratio and factors affecting adult emergence. Ann.
Entomol. Soc. Am. 64: 912-919.
Hoelscher, C.E., R.L. Combs,
fly dispersal. J. Econ.
Jr and J.R. Brazzel. 1968.
Entomol. 61: 370-373.
Horn
Hogsette, J.A. and P.G. Koehler. 19
of horn flies, house flies and
in Florida. Livestock Product
86. Biology and control
stable flies on cattle
ion Pointers 18, 11pp.
Hogsette, J.A., D.L. Prichard and J
effects of horn fly (Diptera:
beef cattle exposed to three p
regimes. J. Econ. Entomol. 84:
.P. Ruff. 1991. Economic
Muscidae) populations on
esticide treatment
1270-1274.


64
pastures year round and were abundant from July to October.
Brontaea spp. were collected in the late summer and fall,all
year round and were very abundant during the summer and
fall. Ceratopogonidae, and Dolichopodidae were mainly
collected in the fall. Psychodidae and Anisopodidae were
mainly collected in the late-fall and winter. Bibilionidae
were collected only in July 1991 and in July-August 1992.
Larvae of Ravinia lherminieri (Robineau-Desvoidy) have
been reported to be predators of other dung-inhabiting flies
(Pickens 1981) R_^ derelicta was abundant and might play
the same role as R_¡_ lherminieri in reducing horn flies.
Sphaeroceridae, Sepsidae Scatopsidae, and Psychodidae are
primarily dung feeders (Skidmore 1991; Valiela 1974).
Sphaerocerid flies are reportedly phoretic on scarab beetles
in Florida (Sivinski 1983). Cecidomyiidae and Sciaridae are
primary fungus feeders, but the larvae of Lestodiplosis sp.
are predacious (Gagn, pers. comm.). The effect of these
larvae on horn fly and other fly pppulations has not been
measured. Empididae and Dolichopodidae have also been
mentioned as predators (Hammer 1941), but their effect on
horn and other flies in the dung is unknown. The larvae of
Bibionidae are big and active in the dung and seem to be
predators of flies, but they are reported to invade the dung
from surrounding soil (Skidmore 19^1). Chloropidae,
Ephydridae, Drosophilidae, Tipulidae and Tabanidae occur
casually in cow-dung. According to Skidmore (1991), the


Table 5-5. Numbers of staphylinid
pitfall traps from the Amdro--
in October 1993.
specimens collected by
treated and control area
126
Subfamily
Species Name
Amdro
Control
Oxytelinae
Anotylus nanus
A. insignitus
Oxytelus incisus
Staphylininae P. flavolimbatus
P. ventralis
Xantholininae Neohypnus pusillus
Aleocharinae
Acrotona hebeticorn.
Atheta sp.
Tinotus brunnipes
T. amplus
Aleochara notula
Others
Tachyporinae M. flavicollis
Total
is
1
44
30
44
0
173
2
5
19
45
238
0
604
2
13
11
3
2
0
75
1
9
4
19
27
168


3
arthropods; 3) to determine the effect of the whole
arthropod community on horn fly production from cattle dung;
4) to determine the species of predatory beetles in cattle
dung and evaluate their predation ability on horn flies; 5)
to determine the effect of the red imported fire ant (RIFA)
on horn fly production in pastures; and 6) to study the
biology of the predatory Staphylinidae found in cattle dung.


30
yielded by scarabs results in potential savings of some two
billion dollars in the USA (Fincher 1981).
The effect of dung beetles on suppression of horn flies
has been best shown in Africa, where upward of 2,000 species
of coprid beetles are known to use the dung of many species
of herbivorous vertebrates (Waterhouse 1974). Therefore,
the horn fly is not a problem to the African cattle
industry. In Britain, Skidmore (1991) reported that
Aphodius contaminatus descend on a cowpat and disrupt other
arthropod colonists. Sometimes A. contaminatus is so
abundant that there may be more beetles than dung and the
effects of such visits are to scatter the dung over a wide
area and render it useless for other community members.
Using coprophagous scarabs to reduce populations of
dung-inhabiting arthropods has received much attention
(Anderson and Loomis 1978; Blume et al. 1973; Bornemissza
1970, 1976; Fincher 1981, 1986). There have been active
programs for the importation and establishment of exotic
coprophagous scarabs in Australia (Bornemissza 1976) and the
United States (Blume et al. 1973).
Although more than 100 species of dung beetles have
been reported from cattle dung on pastures in the U. S.,
millions of cattle droppings remain on the surface of
pastures for several months (Fincher 1990). This is
because native dung beetles and other coprophagous organisms
cannot effectively consume and remove them (Fincher 1986,



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CHAPTER 6
CONCLUSION AND GENERAL DISCUSSION
The Arthropod Commun
itv Fauna
The complete arthropod communi
pats in pastures in north-central F
for the first time. Over 60,000 ar
and identified to over 220 species
in 14 orders. Coleptera contained
Diptera the second (35) and Hymenop
Most of the beneficial insects are
(Harris and Blume 1986).
The fauna of dung-inhabiting a
central Florida is more diverse tha
elsewhere in North America. The ar
includes 226 species in this study,
species reported in Illinois (Mohr
California (Poorbaugh et al. 1968),
(Sanders and Dobson 1966) 109 spec
1969), 103 species in Texas (Blume
Minnesota (Cervenka and Moon 1991),
(Wingo et al. 1974), and 67 species
Canada (Macqueen and Beirne 1975).
ty associated with cattle
lorida was investigated
thropods were collected
belonging to 73 families
most species (109),
tera the third (23).
in these three orders
rthropods in north
n that reported from
thropod community
contrasted with 67
1943), 151 species in
38 species in Indiana
ies in New York (Valiela
1970), 108 species in
157 species in Missouri
in British Columbia,
137


159
Legner, E.F. 1986. The requirement
interactions among dung beetle
natural enemies. pp. 120-131.
A.D. Rutz (eds.): Biological C
Entomol. Soc. Am. Mise. Publ.
for reassessment of
s, symbovine flies, and
In R.S. Patterson and
ontrol of Muscoid Flies.
61, College Park, MD.
Legner, E.F. and I. Moore. 1977. Th
spiculus Erichson (Coleptera:
occurrence in bovine feces in
Psyche 87: 159-164.
e larva of Platvstethus
Staphylinidae) and its
irrigated pastures.
Legner, E.F. and R.W. Warkentin. 19
the dynamics of Onthophagus ga
Scarabaeidae) with symbovine f
Colorado desert of California.
Vector. Control. Assoc. 51: 9-
83. Questions concerning
zella (Coleptera:
lies in the lower
Proc. Calif. Mosq.
11.
Lemke, L.A. and J.B. Kissam. 1988.
fire ant (Hymenoptera: Formici
flies (Diptera: Muscidae) in a
with Pro-Drone. J. Econ. Entom
Impact of red imported
dae) predation on horn
cattle pasture treated
ol. 81: 855-858.
Lindquist, A.E. 1936. Parasites of
Entomol. 29: 1154-1158.
Lofgren, C.S., F.J. Bartlett, C.E.
1964. Imported fire ant toxic
tests with granulated Mirex-s<
Entomol. 57: 695-698.
the horn fly. J. Econ.
Stringer, and W.A. Banks,
bait studies: further
oybean old bait. J. Econ.
Lyle, C. and
ant. J.
I. Fortune. 1948. Notes on an imported fire
Econ. Entomol. 41: 833-834.
Lysyk, T.J. 1992a. Simulating devel
flies, Haematobia irritans (L.
Alberta. Can. Entomol. 124: 84
opment of immature horn
) (Diptera: Muscidae),
1-851.
in
Lysyk, T.J. 1992b. Effect of larval
maternal photoperoid on diapau
(Diptera: Muscidae). Environ.
rearing temperature and
se in the horn fly
Entomol. 29: 1056-1059.
Mackley, J.W., D.A. Carlson and J.F
Identification of the cuticula
horn fly and assays for attrac
. Butler. 1981.
r hydrocarbons of the
tion. J. Chem. Ecol.
660-683 .
7:
Macqueen, A. 1975. Dung as an insec
beetles as competitors of othe
food source: Dung
coprophagous fauna and
as targets for predators. J. Appl. Ecol. 12: 821-827.


167
Tawfik, M.F.S., K.T. Awadallah, E.E
Ela. 1976c. On the bionomics
loncficornis Steph. (Coleptera
Soc. Entomol. Egypte 60: 379-3
Thomas, G.D. 1981. Insect parasites
fly in Missouri, pp. 118-123
(eds.): Status of Biological C
USDA Sci. and Education Admini
Thomas, G.D. and C.E. Morgan. 1972a
fly in Missouri. J. Econ. Ente
Thomas, G.D. and C.E. Morgan. 1972b
of the immature stages of the
Environ. Entomol. 1: 453-459.
Thomas, G.D., I.L. Berry, and C.E.
developmental time of nondiapa
Missouri. Environ. Entomol. 3:
Thomas, G.D., R.D. Hall and I.L. Be
the horn fly (Diptera: Muscida
Entomol. 16: 1092-1097.
Townes, H. and M. Townes. 1962. Ich
north of Mexico. 3. Subfamily
Mesostenini. Smithsonian Inst.
Tugwell, P., E.C. Burns and J.W. Tu
breedings as a factor affectin
repellency of cattle to the ho
62: 56-57.
Tugwell, P., E.C. Burns and B. With
the flight behavior of the hor
irritans (L.) (Diptera: Muscid
Soc. 39: 561-565.
Valiela, I. 1969a. An experimental
factors of larval Musca autumn
Monogr. 39: 119-225.
Valiela, I. 1969b. The arthropod fa
central New York and sources
una of bovine dung in
n its natural history. J.
New York. Entomol. Soc. 77: 20-220.
Valiela, I. 1974. Composition, food
limitation in dung arthropod
invasion and succession. Am. M
Ammar and S.M. Abul-
of Philonthus
Staphylinidae). Bull.
87 .
of the horn fly and face
In R.S. Patterson et al.
ontrol of Filth Flies,
st. Gainesville, Fla.
Parasites of the horn
mol. 65: 169-174.
Field-mortality studies
horn fly in Missouri.
Morgan. 1974. Field
using horn flies in
151-155.
rry. 1987. Diapause of
e) in the field. Environ.
neumon-flies of America
Gelinae, Tribe
Washington, D.C. 602 pp.
rner. 1969. Brahman
g the attractiveness or
rn fly. J. Econ. Entomol.
erspoon. 1966. Notes on
n fly, Haematobia
ae). J. Kans. Entomol.
study of the mortality
alis DeGeer. Ecol.
webs and population
ommunities during
idland Nat. 92: 370-385


24
1990) and members of this family have been reported to be
predators of developing dipterous larvae in cowpats
(Bornemissza 1968; Thomas and Morgan 1972b; Summerlin et al.
1981, 1984c, 1990; Wang et al. 1990). Laboratory tests
(Summerlin et al. 1982, 1984c, 1990) have indicated that
Hister coenosus Erichson is the most effective histerid
species in reducing horn fly populations. It reduces horn
fly populations by 99.1%, when the adult beetles are exposed
to all immature stages of the horn fly. Next in order of
effectiveness are Hister incertus Marseul (98.7%), H.
abbreviatus F. (96%), Atholus rothkirchi Bickhardt (55.8%),
Saprinus pennsvlvanicus (Paykull) (40.9%) and Xerosaprinus
orbiculatus (Marseul) (40.8%). Predation rates increase as
the density of predators increases (Summerlin et al. 1982,
1990). In Texas, Summerlin et al. (1989, 1991) also found
Phelister panamensis LeConte, P. haemorrhous Marseul, and
Pachvlister caffer Erichson to be predators of horn fly
immatures.
Red imported fire ant. Other species of ants, such as
Pocfonomvrmex californicus (Buckley) (Wharton 1979) are
reported as very important predators of horn flies, but the
most emphasizes the red imported fire ant (RIFA), Solenopsis
invicta Burn. This ant arrived near Mobile, Alabama,
approximately 50 years ago from South America and has spread
across the southern United States (Porter and Savignano
1990). RIFA has been labelled a serious agricultural pest


87
(P > 0.05). Most Philonthus adults
3 lived 2 to 3 months
under laboratory conditions and th<
sy consumed numerous horn
fly immatures during their lives.
Larvae of all five species pr<
ayed on horn fly eggs and
larvae. P. lonaicornis had the hial
lest rate, followed by P_j_
ventralis; P. heoaticus had the loi
-zest rate. P¡_
lonaicornis preferred eaas over la]
:vae, but P. ventralis
preferred larvae over eggs. The o'
aher three species preyed
on a slightly higher number of fly
larvae than of eggs
(Table 4-1). The larvae of all f
Lve Philonthus species
began to prey on horn fly eggs or |arvae soon after they
hatched. Predation rates then inc]
reased, peaking on the 6-
7th dav for P. ventralis, and the
1th or 5th day for the
remainina species. P. lonaicornis
preyed on similar numbers
of fly eggs and larvae on the firs'
t day, but on more larvae
than eggs on the following days; P_
flavolimbatus preved
primarily on larvae on the first t\
to days, on more eggs
than larvae on the 3-5th day, and
:hen on more larvae than
eaas; P. sericans preved primarily
t on larvae for the first
three days after hatching, then on
more eggs than larvae
during the following days; and EL_
hepaticus preved onlv on
fly larvae for the first two days,
on a few eggs on the 3rd
and 4th day, and then on more eggs
than larvae (Fig. 4-6).
All the larvae stopped feeding 1-2
days before they pupated.
When five newly hatched larva
2 of P. lonaicornis were
confined individually in Petri dishes and provided manure


28
(Poorbaugh et al. 1968).
they are not disposed of,
Competitors
A horn fly larva needs approximately 2 mg dung to
complete its development (Macqueen and Beirne 1975). A
single adult bovine drops an average of 12 dung pats every
day (Waterhouse 1974), yielding a total weight of some 30
kilograms of feces. This amount of manure can support
15,000 flies (horn and stable flies are the main pests of
cattle in the USA). Moreover, if
the pats produced by each animal will blanket between 5 and
10 percent of an acre in a year. In addition, at the
periphery of each dung pat, there develops a zone of tall,
rank herbage that cattle seldom eat and avoid for a year or
more unless they are ravenous. The effective area of
pasture is thereby reduced by each
of an acre per year (Waterhouse 19
approximately 10.5 million cattle
(Campbell 1993), producing more th
Without competitors, accumulated m
trillions of pest flies and cause
pasture.
Coprophactous beetles. Dung b
long been regarded as useful agent
that develop in dung of domestic a
Lindquist 1936). Since the horn fly lays its eggs in fresh
bovine by about one-fifth
74). There are
in the United States
an 120 million pats a day.
anure pats produce
loss of large areas of
eetles (Scarabaeidae) have
s in the control of flies
nimals (Fullaway 1921;


100
size (8-9 mm long) and
fly immatures, but
of this species were not
but each (n = 5) consumed
fly larvae (range 3-9)
Paederinae
Achenomorphus corticinus. Adults of this species were
observed to prey on horn fly eggs and larvae. Each adult (n
= 6) consumed an average of 10.0 8.55 (SD) eggs (range 0-
26) and an average of 17.0 4.86 (SD) fly larvae (range 9-
24) per day. Adults are of medium-£
have a high predation rate on horn
populations in pastures are low.
Lithocharis sororcula. Adults
observed to prey on horn fly eggs,
an average of 5.4 1.96 (SD) horn
per day. This species was commonly collected in the field,
and adults might be effective predators of horn fly
immatures.
Rugilus ancrularis. Adults of this species were observed
to prey primarily on horn fly larvae (only one fly egg was
observed to be damaged). An average of 12.9 5.01 fly
larvae (range 7-20) was consumed by
per day (n = 8).
These paederine rove-beetle adi
predation potential than do those o:
immatures.
each adult R^_ angularis
ults have higher
f Oxytelinae on horn fly
Xantholininae
Phacophallus tricolor. When t
individually in Petri dishes and we
wo adults were confined
re provided horn fly eggs


166
Summerlin, J.W., D.E. Bay, K.C. Sta
III. 1984c. Laboratory observa
and habits of Hister abbreviat
fford and J.S. Hunter
tions on the life cycle
us (Coleptera:
Histeridae) Ann. Entomol. Socj. Am. 77: 543-547.
Summerlin, J.W., G.T. Fincher and J
by Atholus rothkirchi on horn
15: 253-256.
Summerlin, J.W., G.T. Fincher, J.P.
.P. Roth. 1990. Predation
fly. Southwest. Entomol.
Roth and H.D. Petersen.
1989. Laboratory studies of the life cycle and prey
relationships of Pachvlister c.
(Coleptera: Histeridae). J. E
Summerlin, J.W., R.L. Harris and H.
important fire ant (Hymenopter
and intensity of invasion of f
Environ. Entomol. 13: 1161-116
Summerlin, J.W. and S.E. Kunz. 1978
imported fire ant on stable fl
3: 260-262.
Summerlin, J.W., S.M. Moola, G.T. F
1991. Laboratory observations
Phelister panamensis LeConte
including scanning electron mi
stages. J. Agrie. Entomol. 8:
incher and J.P. Roth,
on the life cycle of
(Coleptera: Histeridae)
croscopy of the life
189-197.
Summerlin, J.W., J.K. Olson, R.R. B
Bay. 1977. Red imported fire
Onthophagus gazella and the ho
6: 440-442.
Summerlin, J.W., H.D. Petersen and
imported fire ant (Hymenoptera
the horn fly (Diptera: Muscida
scarabs. Environ. Entomol. 13:
Tawfik, M.F.S., K.T. Awadallah, E.D
Ela. 1976a. The life history
(Coleptera: Staphylinidae). B
60: 345-355.
Tawfik, M.F.S., K.T. Awadallah, E.E
Ela. 1976b. The life history
Philonthus turbidus Er. (Cole
Bull. Soc. Entomol. Egypte 60:
affer Erichson.
ntomol. Sci. 24: 329-338.
D. Petersen. 1984a. Red
a: Formicidae): frequency
resh cattle droppings.
3 .
. Predation of the red
ies. Southwest. Entomol.
lume, A. Aga and D.E.
ant: effects on
rn fly. Environ. Entomol,
R.L. Harris. 1984b. Red
Formicidae): effects on
e) and coprophagous
1405-1410.
Ammar and S.M. Abul-
of Philonthus misor Tott.
ull. Soc. Entomol. Egypte
Ammar and S.M. Abul-
of the staphylinid
qptera: Staphylinidae) ,
357-366.


74
community, especially predators, on horn flies.
Objectives of the following study were to determine
horn fly mortality caused by the whole arthropod community
in pastures, and to evaluate the predation potential of
individual predatory species on horn flies.
Materials and Methods
Tests of the Arthropod Community on Horn Fly Survivorship
Laboratory studies on horn fly survivorship
Survivorship of horn flies was studied in the
laboratory before initiating field studies. The horn flies
used were from the colony maintained by Dr. J. F. Butler,
Department of Entomology/Nematology, University of Florida.
Standard colony rearing methods were developed by Greer
(1975) and modified by Okine and Butler (1995). Larval
medium was the mixture of frozen and thawed manure and
peanut hulls. The adults were fed with bovine blood. The
colony was maintained in an environmental chamber at 27
3C and 75 5% RH with continuous light.
Manure used for experiments was collected in a pasture
within 30 min after its deposition, frozen for 48 hours and
thawed for 24 hours. The tests were conducted at 27C, 50-
60% RH and 14:10 (L:D) photoperiod.
Artificial pats were prepared by following the


65
larvae of most chloropid species develop in the stems of
grasses; the larvae of Ephydridae develop in a range of
materials, including muddy soil, rotting vegetation, or cow-
dung; and Drosophilidae develop mostly in decaying plant or
animal matter in which fermentation is taking place.
Drosophilidae were abundantly seen in decaying mushrooms
growing in pastures in the present study. Tipulid larvae
occur in dung, but belong to the soil fauna.
Hvmenoptera
A total of 24 species belonging to 15 families of
Hymenoptera was collected. The family with the most
represented species was Formicidae, of which the red
imported fire ant (RIFA), Solenopsls invicta, was abundant
all year round and peaked in September-October. RIFA has
been well documented as a predator of flies (Laurence 1954;
Bruce 1964), in particular, of horn flies (Howard and Oliver
1978; Summerlin et al. 1984b; Schmidt 1984). The effect of
RIFA on the horn fly will be discussed in chapter 5.
Two species of eulophids were collected: Aprostocetus
spp. have been recorded to be parasitoids of gall-midges,
and Trichospilus spp. as parasitoids of fly puparia
(Krombein et al. 1979). Among the Braconidae collected, one
species (Aphidiinae) is a parasitoid of aphids. The other,
Aphaereta pallipes. was commonly collected by trapping and
rearing from extracted sarcophagid puparia; its hosts


152
Figg
D.E., R.D. Hall, and G.D. Tho
parasites associated with Dipt
dung pats on central Missouri
Entomol. 12: 961-966.
mas. 1983. Insect
era developing in bovine
pastures. Environ.
Fincher, G.T.
pasture
1981. The potential value of dung beetles in
ecosystems. J. Ga. Entomol. Soc. 16: 316-333.
Fincher, G.T. 1986. Importation, cc
of dung-burying scarabs, pp. 6
and D.A. Rutz (eds.): Biologic
Flies. Mise. Pub. Entomol. Soc
Ionization, and release
9-76. In Patterson, R.S.
al Control of Muscoid
. Am. 61, College Park,
MD.
Fincher, G.T. 1990. Biological cont
flies: Pest of pastured cattle
pp. 137-151. In Rutz, D.A. and
Biocontrol of Arthropods Affec
Poultry, Westview Press, Bould
rol of dung-breeding
in the United States.
R.S. Patterson (eds.):
ting Livestock and
er, Colorado.
Fincher, G.T. 1992. Injectable ivermectin for cattle:
Effects on some dung inhabiting insects. Environ.
Entomol. 21: 871-876.
Fincher, G.T. and Morgan. 1990. Fli
and poultry, pp. 145-155. In D
and J.H. Frank (eds.): Classic
the Southern United States. So
Series, 315. Gainesville, FL.
es affecting livestock
.A. Habeck, F.D. Bennett
al Biological Control in
uthern Cooperative
/
Fincher, G.T., T.B. Stewart, and J
S. Hunter, III. 1983. The
1981 distribution of Onthophagiis qazella Fabricius from
releases in Texas and Onthophagus taurus Schreber from
an unknown release in Florida
Scarabaeidae). Coleopt. Bull.
(Coleptera:
37: 159-163.
Fincher, G.T. and J.W. Summerlin. 1994. Predation on the
horn fly by three exotic species of Philonthus. J.
Agrie. Entomol. 11: 45-48.
Fincher G.T. and R.E. Woodruff. 197
beetle, Onthophagus taurus Sch
(Coleptera: Scarabaeidae). Co
5. A European dung
reber, new to the U.S.
leopt. Bull. 29: 349-350.
Foil
L.D., C.S. Foil, D.D. French,
Miller. 1990. The role of horn
management of seasonal equine
dermatitis. Equine Practice 1
T.R. Klei and R.I.
fly feeding and the
ventral midline
2: 6-14.
Foil, L.D. and J.A. Hogsette. 1994. Biology and control of
tabanids, stable flies and horn flies. Rev. sci. tech.
Off. int. Epiz. 13: 1125-1158.


107
Though staphylinid species hav
re been considered
effective predators of the flies, p
revious evidence was
provided for predation on horn fly
immatures only for
Philonthus and Aleochara species.
The present study gives
evidence of predation on horn fly e
ggs and larvae by other
genera and species.
Hvdrophilidae
Cercvon varieaatus. When adul
t C. varieaatus were
confined individually in Petri disi
es and provided horn fly
eggs and larvae for food, without it
anure, no horn fly eggs
and larvae were observed to be dama
ged and killed. All
varieaatus starved to death Cn = 5}
The result confirms
earlier findinas that Cercvon sod.
are generally
coprophagous (Sanders and Dobson 15
66; Hafez 1939; Merritt
1976; Thomas and Morgan 1972b).
Another commonly collected spe
cies of Hydrophilidae in
the survey was Sphaeridium lunatum
which was previously
reported as an effective predator c
f horn flies (Macqueen
and Beirne 1975). The adults of tf
iis species were observed
to prey on horn fly eggs and larvae
., but no tests were
conducted on them.
Histeridae
Hister coenosus was reported a
is one of most effective
predatory species of histerid beet!
.es (Summerlin et al.


18
coleopterous and hymenopterous species. Information from
North America shows that arthropod species that affect horn
flies vary geographically, but overall groups are generally
the same (Harris and Blume 1986). On the other hand,
information from South Africa shows that the dung faunal
species composition and abundance differ between habitats,
although a degree of overlap exists (Fay 1986). Each
habitat has its own distinct community of dung organisms
that are effective control agents of horn flies. There is
only a small core of common species that occur in all the
habitats and these play the major role in maintaining the
fly population at a certain level (Fay 1986). Most of the
beneficial insects belong to the orders Coleptera, Diptera
and Hymenoptera (Fincher 1990; Harris and Blume 1986) with
many other arthropods of less importance (Sanders and Dobson
1966). Mohr (1943) was one of the first workers in the
United States to consider the entire insect complex of
cattle droppings. Since then, several comprehensive surveys
have been conducted throughout the country (Macqueen and
Beirne 1975; Merritt and Anderson 1977; Sanders and Dobson
1966; Poorbaugh et al. 1968; Blume 1970; Valiela 1969b;
Wingo et al. 1974). Studies on specific groups of the
arthropods in pastures in America north of Mexico were
summarized by Blume (1985), who included a checklist,
distribution maps, and an annotated bibliography. Over 400
species of arthropods have been collected in or on cattle


45
Table 3-2. Staphylinidae collected in pastures in north
central Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and
(+++) abundant.
Trapped
Extracted
Osorinae
Osorius sp.
1
Oxytelinae
Oxytelus incisus Motschulsky
617
2365
Platystethus spiculus Erichson
0
57
P. americanus Erichson
14
109
Anotylus nanus (Erichson)
65
79
A. insignitus (Gravenhorst)
129
67
Apocellus sphaericollis (Say)
15
26
Paederinae
Rugilus angularis (Erichson)
26
22
Achenomorphus corticinus
2
+
(Gravenhorst)
Lithocharis sororcula (Kraatz)
27
34
Thinocharis sp.
10
36
Homaeotarsus cinctus (Say)
2
+
Astenus fusciceps (Casey)
3
+
Scopaeus sp.
1
+
Staphylininae
Philonthus hepaticus Erichson
11
107
P. flavolimbatus Erichson
95
53
P. longicornis Stephens
91
38
P. rectangulus Sharp
2
+?
P. sericans (Gravenhorst)
5
26
P. ventralis (Gravenhorst)
16
36
Endius sp.
2
Gabronthus mgogoricus Tottenham
1
Platydracus tomentosus
4
0
(Gravenhorst)
Xantholininae
Phacophallus tricolor (Kraatz)
3
+
Lithocharodes ruficollis
8
+
(LeConte)
Neohypnus pusillus (Sachse)
15
119
N. attenuatus (Erichson)
6
32
N. emmesus (Gravenhorst)
2
11


140
The fauna-caused horn fly mortality in pastures in the
present study was lower than those reported previously, such
as 87.9% in Texas (Roth 1989), and 97.7% in Missouri (Thomas
and Morgan 1972b).
Mortality of horn flies caused by the dung arthropod
fauna in pastures is significant. The beneficial insects
associated with cattle dung are diverse and numerous. Dung
beetles compete with the horn fly f
or food, interrupt the
habitat of the immature flies, and
reduce dung accumulation
on pasture. Beetles in the familie
s Staphylinidae,
Hydrophilidae, Histeridae, and Cara
bidae are predators and
contribute to the control of the ho
rn fly. Larvae of many
fly species develop in the pat and
compete for food or prey
on immature horn flies. In addition
to these natural
enemies, hymenopterous parasites an
d ants further reduce fly
populations. However, the numbers
of horn flies which
survived over predation, competitio
n, and parasitization by
natural enemies reach or exceed the
economic threshold to
the cattle industry at some times a
nd places in the USA.
The reasons of the failure to
control the horn fly by
the arthropod fauna may be: 1) the
inadequate insect fauna
diversity compared with its origin-
-Europe; 2) the lack of
host- and habitat-specific predator
s and parasitoids; 3)
scarab beetles do not function well
on burial and shredding
of dung; 4) immature horn flies may
survive in the dung
buried by the beetles because of th
e humid weather in north


158
Kunz, S.E., R.R. Blume, B.F. Hogan
Biological and ecological inve
in central Texas: Influence of
deposition on oviposition. J.
933 .
Kunz, S.E. and J.L. Eschle. 1971. P
sterile male technique for era
pp. 145-156. In Proc. of a Syirv
Principle for Insect control
International Atomic Energy Ag
Kunz, S.E., M.R. Graham, B.F. Hogan
Effect of releases of sterile
population of horn flies. Envi
Kunz, S.E., B.F. Hogan, R.R. Blume,
Some bionomical aspects of hor:
central Texas. Environ. Entorno
Kunz, S.E., K.D. Murrell, G. Lamber
Terrill. 1991. Estimated losse
pp. 69-98. In D. Pimentel (ed.
Management in Agriculture. Vol
Kunz, S.E. and C.D. Schmidt. 1985.
problem in the horn fly. J. Ag:
Laake, E.W. 1946. DDT for the contr
Kansas. J. Econ. Entomol. 39:
Lancaster, J.L. and M.V. Meisch. 19
livestock and poultry producti
Chichester, England, pp. 90-10
Laurence, B.R. 1952. The prey of so
Dolichopodidae (Dipt.). Entom
157.
Laurence, B.R. 1954. The larval inh
Anim. Ecol. 23: 234-260.
and J.J. Matters. 1970.
stigations of horn flies
time of manure
Econ. Entomol. 63: 930-
ossible use of the
dication of the horn fly.
posium on the Sterility
r Eradication,
ency. Vienna, 542 pp.
and J.L. Eschle. 1974.
horn flies into a native
ron. Entomol. 3: 159-162.
and J.L. Eschle. 1972.
n fly populations in
1. 1: 565-568.
t, L.F. James and C.E.
s of livestock to pests,
): CRC Handbook of Pest
. 1, CRC, Coca Raton, FL.
The pyrethroid resistance
ric. Entomol. 2: 356-363.
ol of the horn fly in
966-972.
86. Arthropods in
on. Ellis Harwood Ltd.
6.
me Empididae and
ol. Mon. Mag. 88: 156-
abitants of cow pats. J.
Legner, E.F. 1978. Natural enemies
for the biological control of
autumnalis DeGeer, and horn fl
(L.). Proc. Calif. Mosq. Vecto
79 .
imported in California
face fly, Musca
y, Haematobia irritans
r. Contr. Assoc. 46: 77-


33
the predators for horn fly biocontrol is questionable.
Large numbers of scarabs burrowing through the dung could
cause significant dung removal and pasture improvement
(Fincher 1981; Legner 1986; Waterhouse 1974) but might not
achieve significant fly reductions (Legner 1978, 1986;
Macqueen 1975). High numbers of dung beetles could disrupt
the oviposition behavior of female staphylinids and reduce
the food source of immatures, causing a decrease of the
numbers of predators (Legner and Warkentin 1983; Roth 1983),
hymenopterous parasitoids, and coprophagous Coleptera (Roth
et al. 1991). Therefore more emphasis should be placed on
predators and parasitoids as biological control agents
(Legner 1978; Roth 1983).
Coprophagous flies. The adults and larvae of some
dipteran flies feed on dung, microorganisms (e.g. fungi,
bacteria) growing in the dung, or
matter (e.g., fungal spores and hyphae) (Merritt 1976).
Though some species of Muscidae and Sarcophagidae may
compete with the horn fly in central Texas (Kunz 1978), the
populations of these flies are us|ually not as high as those
of coprophagous beetles, because they encounter predation
and competition. Small dung flies,
very high populations, but their bi
less (Harris and Blume 1986). Macqueen and Beirne (1975)
thought that dung-inhabiting Diptera do not encounter food
limitations and Poorbaugh et al. (1968) regarded competition
decomposing vegetable
, such as Sepsidae, have
iomass is considerably


97
Petri dishes with horn fly eggs and larvae but no cow dung,
feeding on first instar fly larvae occurred, but the beetle
larvae did not pupate. However, when the beetle larvae were
reared with cow dung and horn fly eggs and larvae, they
pupated successfully in chambers or partial chambers under
dung.
The food range of Platystethus adults and larvae has
(1991), without providing
been studied previously. Mohr (1943) assumed that P.
americanus adults prey on fly larvae, because it is a
staphylinid, and Cervenka and Moon
evidence, considered it to be predatory. However, the
family Staphylinidae includes fungivores as well as
predators. Platystethus arenarius (Fourcroy) adults and
larvae were observed by Hinton (1944) to ingest cattle dung,
and were stated by Skidmore (1991)
1,
to feed exclusively on
cow-dung. Larvae of P. spiculus were reported from cow dung
by Legner and Moore (1977). Frank (1976) found adult P.
spiculus in horse dung, but many were discovered feeding on
a slime mold [Fuligo sptica (L.) Wigger]. Palomino and Dale
(1989) reported collecting P_¡_ spiculus adults from cattle
dung, obtaining eggs, and rearing larvae on a diet of house
fly eggs (killed in hot water) in small Petri dishes.
Feeding on horn fly eggs and larvae was first reported in
the present study.
Anotvlus insignitus. Adults (n = 26) of this species
were not observed to prey on horn fly eggs, but consumed an


39
composition and abundance of the arthropod community
associated with cattle dung, and to monitor seasonal
distribution and diel activity of arthropods found.
Materials and Methods
A survey of the arthropod community associated with
1991 and from June 1992 to
September-December 1993
is 16 km (10 miles)
cattle dung was conducted in July
December 1993 in pasture A and in
and 1994 in pasture B. Pasture A
northeast of Gainesville, Alachua County, Florida, and
contained approximately 250 beef cattle at the time of the
study. Pasture B is on the southern side of Gainesville and
contained 40 cattle at the time of the study. Two methods
of collection were used: pitfall traps baited with fresh
cattle dung, and emergence boxes (Fig. 3-1) that held entire
cowpats and trapped all emerging arthropods. A dilute soap
solution was added to the pitfall
arthropods and keep them clean and flexible for subsequent
processing. Ten to 15 pitfall traps were set twice each
month from May to October, and once each month from November
to April in pasture A; 20 traps were set in September and
October 1992 and 1993 in pasture B, respectively.
Arthropods captured in the traps were collected after 24
hours and preserved in 70% alcohol for identification.
traps to drown trapped
Between December 1992 and December 1993, cattle dung


148
Bourne, J.R. and K.L. Hays. 1968.
predation of horn fly larvae
Sohaeridium scarabaeoides. J.
322 .
ffect of temperature on
by the larvae of
Econ. Entomol. 61: 321-
Bourne, J.R. and K.L. Hays. 1973. Predators of the horn fly
in the Piedmont of Alabama. Highlights Agrie. Res. 20:
886.
Bourne, J.R. and R.J. Nielsson. 19^7. Nemobius fasciatus A
predator on horn fly pupae. J. Econ. Entomol. 60: 272-
274 .
Brown, Y.W. 1940. Notes on the Amer
species of Coleptera common
America continents. Can. Entor
ican distribution of some
o the European and North
ol. 72: 65-78.
Bruce, W.G. 1938. A practical trap
flies in cattle. J. Kansas Ent
for the control of horn
omol. Soc. 11: 88-93.
Bruce, W.G. 1942. The horn fly, pp.
(ed.): Keeping Livestock Healt
Yearbook of Agrie., Washington
626-630. In G.
hy. U. S. Dept.
, D.C. 1276 pp.
Hambridge
Agrie.
Bruce, W.G. 1964. The history and b
Haematobia irritans (Linnaeus)
control. N.C. Agrie. Exp. Stn.
i
r
ology of the horn fly
with comments on
Tech. Bull. 157: 1-33
/
Bruce, W.G.and G.C. Decker. 1947.
J. Econ. Entomol. 40: 530.
iy
control and milk flow.
Butler, J.F. 1975. Economics and co
insects, pp. 143-152. In Proc.
Training Manual-Radio Isotopes
Entomology, Gainesville, Fla.
ntrol of livestock
Short Course FAO/IAEA
and Radiation in
Butler, J.F. 1990. Haematobia irritans: Economic importance
and bionomical characteristics. XVI Congresso Mundial
de Buiatria, VI Congresso Latino Americano de Buiatria.
Butler, J.F., R.L. Escher and J.A.
parasite levels in house flies
flies in Florida, pp.61-79. In
Status of Biological Control o
Proceedings of a Workshop. U.S
Education Administ. Gainesvill
logsette. 1981. Natural
, stable flies, and horn
R. S. Patterson (ed.):
f Filth Flies.
. Dept. Agrie. Sci.
5, Fla.
Butler, J.F., W.J. Kloft, L.A. Debose and E
Recontamination of food after feeding
biting Muscidae. J. Med. Entomol. 13:
.S. Kloft. 1977.
a 32p source to
567-571.


32
Bornemissza (1960) advocated the use of certain species
of dung beetles in Australia to effect disposal of surface
dung, improve pastures by incorporation of dung into soils,
and reduce larval habitats for the horn fly. The CSIRO
Division of Entomology began a program to import foreign
dung-burying scarab beetles into Australia in 1964
(Bornemissza 1976). The purpose of the project was to reduce
the populations of the horn fly in north Australia
(Waterhouse 1974; Bornemissza 1976) by increasing the
natural dispersal of cattle dung. By 1991, CSIRO had
imported and released 55 species qf dung beetles in
Australia (Doube and Macqueen 1991). Colonized scarab
species cause high mortality of horn flies and other
dipterans when their populations are high (Roth et al.
1988a, 1991).
Many species of Hydrophilidae and Staphylinidae, which
are not usually referred to as dung beetles, also use some
components of the decomposing material or the microorganisms
in dung pats (Hanski 1991). Whether they suppress the horn
fly production has not been specifically reported, but
Merritt (1976) and Valiela (1969b, 1974) reported that
Sphaeridium burrowed in and out of the dung shortly after
pats were dropped, providing aerat:
permitting staphylinids and parasil
their tunnels to locate their targets.
On the other hand, the compatibility of scarabs with
ion to the pat and
tic Hymenoptera to use


154
Hammer, O. 1941. Biological and ecc
flies associated with pasturin
excrement. Vidensk. Medd. Natu
141-393.
Hanski, I. 1991. The dung insect co
Hanski and Y. Cambefort (eds.)
Princeton University Press, Pr
371 pp.
Hanski, I. and P. Hammond. 1986. As
dung Staphylinidae in tropical
Borneo. Annales Entomologici F
Hargett, L.R. and R.L. Goulding. 19
Haematobia irritans (L.). J. E
566.
Hargett, L.R. and R.L. Goulding. 19
logical investigations of
g cattle and their
rhist. Foren. Kbh. 105:
mmunity. pp 5-21. In I,
: Dung Beetle Ecology,
inceton, New Jersey,
semblages of carrion and
rain forests in Sarawak,
ennici 52: 1-19.
62a. Rearing the horn fly
con. Entomol. 55: 565-
62b. Studies on the
behavior of the horn fly, Haematobia irritans (Linn.).
Oregon Agrie. Exp. Stn. Tech. Bull. 61. 27 pp.
parasites, predators,
£>ulations. pp. 125-127.
s of biological control
kshop. U.S. Dept. Agrie,
sville, FL.
Beneficial insects
the United Stages, pp.
atterson (eds.):
ntomol. Soc. Am. Mise.
Harris, R.L. 1981. The influence of
and competitors on horn fly po
In R.S. Patterson (ed.): Statu
of filth flies. Proc. of a Wor
Sci. Education Administ. Gaine
Harris, R.L. and R.R. Blume. 1986.
inhabiting bovine droppings in
10-15. In D.A. Rutz and R.S. P
Biocontrol of Muscoid Flies. E
Publ. No. 61, College Park, MD
Harris, W.G. and E.C. Burns. 1972.
star tick by the imported fire
1:362-365.
Harris, R.L., E.D. Frazar and C.D.
the mating habits of the horn
61: 1639-1640.
Harris, R.L., J.A. Miller and E.D.
and stable flies: Feeding acti
Am. 67: 891-894.
Harris, R.L. and L.M. Oliver. 1979.
flavolimbatus on the horn fly.
259-260.
Predation on the lone
ants. Environ. Entomol.
Schmidt. 1968. Notes on
fly. J. Econ. Entomol.
Frazar. 1974. Horn flies
vity. Ann. Entomol. Soc.
Predation of Philonthus
Environ. Entomol. 8:


35
emerges as an adult beetle. The adult beetles are
predacious on eggs and larvae of muscoid Dptera (Wingo et
al. 1974, Klimaszewski 1984). A report (Escher 1977) that
Tinotus planulus Notman and Tinotus sp. are parasitoids has
not been substantiated; and the report that Oxvtelus incisus
is a parasitoid is certainly erroneous.
The level of parasitoidism of the horn flies can be as
high as 43% (Combs and Hoelscher 1969) or 45% (Harris and
Summerlin 1984), but usually it is low and only occasionally
is the level adequate to reduce horn fly populations
significantly (Harris and Blume 1986; Macqueen and Beirne
1975). Wharton (1979) collected thousands of horn fly pupae
from California in spring and summer, but no parasitoids
were found.
Ten hymenopterous species parasitoidal on the horn fly
pupae in Florida were reported by Escher (1977) and Butler
et al. (1981), with an overall level of parasitoidism of
10.5% (from 1.9 in April to 17.7 in August).
Ecology and Biology of Staphvlinidae
Specific information on the biology, ecology and
behavior of many species of dung-inhabiting predatory
beetles, especially the Staphylinidae, is scant. However,
this information is very useful in evaluation of native and
exotic species as biological control agents of the horn fly,
in mass propagation of efficient species for initial release


Table 3-7 (continued)
69
Trapped
Extracted
Collembola
Hypogastruridae
Hypogastrura sp.
+++
+++
Xenylla we1 chi Folsom
+++
+++
Neanura muscorum Templeton
+++
+++
Sminthuridae
Sminthurus jupiterensis Snider (new)
++
+++
Bourletiella gibbonsi Snider
+++
++
Entomobryidae
Lepidocyrtus violaceus Fourcroy
++
+++
L. cf. helenae Snider
++
++
L. cyaneus (Tullberg)
++
+++
Tomocerus elongatus Maynard
++
++
Salina wolcotti Folsom
++
++
Salina (sp. new)
+
++
Seira brasiliana (Arl)
+
++
S. brasiliensis Snider
++
++
S. caheni Jacquemart
++
++
Entomobryoides sp.
+
+
Dermaptera
Labiduridae
Labidura riparia (Pallas)
8
+
Thysanoptera (1 species)
+
+
Isoptera (1 species)
1
+
Arachnida (Acari)
675
+++
Anistidae
Prostigmata sp.
+
+
Parasitidae
Parasitus fimetorus (Berlese)
++
++
Parasitus sp.
+
+
Macrochelidae
Glyptholaspis confusa (Foa)
++
++
Glyptholaspis sp.
+
+
G. pamericana Berlese
+
+
G. fimicula (Sellnick)
+
+
Microcheles mammifer Berlese
+
+
Microcheles sp.
+
M. peniculatus Berlese
++
++
M. muscaedomesticae (Scopoli)
+
+
Oribatidae (1 species)
187
++


21
induced mortality of horn flies was from 79% to 84% and
fauna-reduced headcapsule width of the emerged horn flies
was an average of 2% to 7%. Small flies produced by
indirect effect of the dung fauna are less fecund and have
lower survival potential (Fay 198(5; Roth 1989).
Predators
Non-arthropod animals, especially birds, have been
considered predators of flies (Hammer 1941), but research
attention has been focused on arthropods inhabiting cattle
dung. Studies in Missouri (Thomas and Morgan 1972b), Texas
(Blume et al. 1970; Roth 1983, 1989; Roth et al. 1983), and
Canada (Macqueen and Beirne 1975) indicated that predators
are the primary insect biotic mortality factors affecting
the horn fly.
Studies have shown that the most important predators
are coleopterans of the families Staphylinidae,
Hydrophilidae, and Histeridae (Bornemissza 1968; Fay 1986;
s
Fay and Doube 1983; Fincher 1990; Legner 1986; Macqueen and
Beirne 1975; Roth 1989; Roth et al. 1983; Sanders and Dobson
1969; Thomas and Morgan 1972b), especially Staphylinidae
(Hammer 1941; Blume et al. 1970; Macqueen and Beirne 1975;
Thomas and Morgan 1972b).
Staphylinidae. Staphylinidae are the most important
predators because of species diversity and high population,
and predation rate (Fincher 1990; Harris and Blume 1986).


14
disperse 5 km or even further (Sheppard 1994). The movement
occurs nocturnally (Hoelscher et al. 1968) and dispersing
populations are predominantly females (Marley et al. 1991).
Control of the Horn Fly
Chemical Control
Control of horn flies with chemical insecticides has
been the primary method for the last century and it is
widely practiced today. Methods of insecticide application
include sprays, dust bags, ear tags, tail tags, leg bands,
and back rubbers for killing the adults, and feed-through
for killing fly larvae (Beadles et al. 1975; Butler and
Koehler 1979; Drummond et al. 1988; Hogsette and Koehler
1986; Schreiber et al. 1987a). Beginning with the
widespread use of DDT and related chlorinated hydrocarbon
insecticides, and later with organophosphorus compounds,
horn flies were controlled effectively (Morgan and Thomas
1974, 1977). The advent of insecticidal ear tags (Byford et
al. 1985; Harvey and Brethour 1970, 1979; Quisenberry and
Strohbehn 1984; Schmidt and Kunz 1980) permitted highly
effective season-long horn fly control in most regions.
In the USA, insecticide resistance in horn flies
occurred sporadically before the 1980s when horn flies were
easy to control with the available organochlorine and
organophosphate insecticide. Two to three years after the


89
Fig.
CD
Q
T9
CD
E
ID
09
C
O
C)
09
CD
V-
ID
4'
CD
E
E
o
H
140
120
100
80
60
40
20
0
W/
'A
Larvae
Eggs
B
longicornis (46) serie
flavolimbatus (
ans (52) ventralis (10)
4) hepaticus (19)
4-5. Adult predation rates of five species Philonthus
on horn fly eggs and larvae per day under laboratory
conditions. Different letters; of inside labels
indicate significant differences in mean predation
rates (P < 0.05). Numbers following each species
indicate replicates.


THE ARTHROPOD COMMUNITY IN PASTURES AND ITS BIOCONTROL
POTENTIAL FOR THE HORN FLY, Haematobia irritans (L.)
IN NORTH-CENTRAL FLORIDA
By
GUANGYE HU
0
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1995


41
Fig. 3-1. Emergence boxes used for extracting arthropods
from intact cowpats 24 hrs old.


95
No. Eggs Seeded
Fig. 4-8. Predation of P. lonoicornis on horn flies at
different prey densities under simulated field
conditions (100 g. manure: 0, 50, 100, 250 and 500 horn
fly eggs/replicate).


effect on horn flies than competito
143
rs and parasitoids in
north central Florida. Obvious dun
g pat interruption was
only observed in May when the horn
fly population is not
high (Butler pers. comm.). The pre
sent study also shows
that the most diverse and abundant
family is Staphylinidae
and most of its species were effici
ent predators to the horn
fly eggs and larvae. Therefore, th
e fauna induced horn fly
mortality is mainly attributed to p
redators.
In pastures, many species of p
redatory arthropods may
not be obligate predators. Some ma
y alternatively consume
dung and fungus when prey are not a
dequate. Fungus-feeding
and dung-feeding insects have diges
tive enzymes in their
guts required for the digestion of
cellulose (Martin and
Martin 1978). Detection of these e
nzymes should help to
confirm whether they are obligate c
r facultative predators.
Furthermore, most predators in the
dung may prey on whatever
they encounter and are able to hand
le, such as diverse fly
larvae, mites, and collembolans. 1
o confirm whether they
primarily prey on immature horn fli
es in the dung pats,
electrophoretic, radioactive labell
ing, and immunological
methods (such as enzyme-linked immc
nosorbent assay, ELISA)
should be able to detect the remair
s of immature horn flies
in the guts of predators that are e
xtracted from fly-
infested pats in the field (Doube 1
986) .


139
Doube and Moola 1988; Ferrar 1975; Fincher 1981, 1986, 1990;
Waterhouse 1974). The species number of Staphylinidae
collected in this study is about the same as the total
species numbers reported elsewhere in the USA. But species
numbers of Scarabaeidae and Histeridae are smaller than the
total numbers reported in the other parts of the country
(Blume 1985; Fincher 1990; Harris and Blume 1986).
Higher Diptera collected abundantly were members of
Sarcophagidae and Muscidae, and lower Diptera were members
of Cecidomyiidae, Sphaeroceridae, and Sepsidae. The effect
of those dipterans on horn flies is yet poorly documented.
They are generally divided into coprophagous, facultative
predators and obligate predators (Hanski 1991) .
Horn Fly Mortalities Caused by the Arthropod Fauna
The mortality of horn flies c
community was determined by seeding
artificially formed dung pats. The
exposed to or isolated from the ar
cone traps. The number of horn fl
covered by the cone traps was sign
that from uncovered pats in July a
Mortalities of horn flies in the f
arthropod community were 75.9% and
aused by the arthropod
horn fly eggs underneath
se pats were either
thropod community by using
es that emerged from pats
ficantly greater than
nd August 1992.
eld attributed to the
66.7% in July and August,
respectively. The overall average was 71.3%.


LIST OF FIGS
Figure Page
3-1. Emergence boxes used for extracting arthropods from
intact cowpats of 24 hrs old 41
3-2. Seasonal distribution of five families of Coleptera
most commonly collected by trapping 43
3-3. Seasonal distribution of species diversity of
Staphylinidae and Scarabaeidae in pastures of north
central Florida 44
3-4. Seasonal distribution of staphylinid species in
pasture A, collected by trapping 47
3-5. Seasonal distribution of scarabaeid species in pasture
A, collected by trapping 54
4-1. Cone traps used for covering cowpats to collect horn
and other flies emerged from the pats 79
4-2. Cages used for testing predation rates of
predators on horn fly immatures 8 0
4-3. Horn flies emerged from covered and uncovered cowpats
in a north central Florida pasture in 1992 .... 85
4-4. P^_ loncricornis adult preying on horn fly eggs under
laboratory conditions 88
4-5. Adult predation rates of five species Philonthus on
horn fly eggs and larvae per day under laboratory
conditions 89
4-6. Daily predation rates of 5 species Philonthus larvae
on horn fly eggs and larvae under laboratory
conditions 90
4-7. Predation rates by four species of Philonthus on horn
flies under simulated field conditions 94
IX


were found to be slightly damaged,
99
but an average of 1.9
1.46 (SD) larvae were consumed per
beetle per day (range 0-
4). No beetle eggs were found in t
he Petri dishes. When
the beetles were offered horn fly e
ggs and larvae, as well
as cow manure, larvae of 0. incisus
were found in the manure
a few days later. When the larvae
were removed from the
manure, confined individually in F
etri dishes, and provided
horn fly eggs and larvae for food,
without manure, predation
on horn fly larvae (3-5 fly larvae
per beetle larva)
occurred, but they died without pup
ation during the
wandering stage. The results indie
ate that 0. incisus
adults may also need manure as a su
bstrate for oviposition,
and the larvae need manure in which
to make pupal chambers.
Apocellus sphaericollis. Wher
five adults of A^.
sphaericollis were confined individ
lually with horn fly eggs
and larvae, without manure, each be
etle consumed an average
of 2.4 0.49 (SD) fly larvae, but
no eggs.
The results show that members
of the subfamily
Oxytelinae prey on horn fly larvae,
but predation rates were
low. The oxyteline rove-beetles ai
e at least facultative
carnivores (Skidmore 1991) in their
adult and larval stages.
Species of the genera Platvstethus,
Anotvlus. Oxvtelus need
cattle manure either for adults to
oviposit or for larvae to
pupate.


Table 3-6. Hymenoptera collected in pastures in north
central Florida. The occurrence codes for species not
enumerated: (+) rare, (++) common, and (+++) abundant.
Trapped
Extracted
Pompilidae
Anoplius sp.
+
sp. 2 (Genus?)
+
+
Eulophidae
Trichospilus sp.
5
+
Aprostocetus sp.
2
++
Braconidae
Aphaereta pallipes (Say)
17
++
Sp. 2 (Aphidiinae)
+
+
Pteromalidae
Spalangia cameroni Perkins
10
++
Formicidae
Solenopsis invicta Burn
38250
+++
Pheidole dentata Mayr
+
+
P. metallescens Emery
+
+
Cyphomyrmex sp.
+
4-
Pseudomyrmex mexicana
+
+
Hypoponera opaciceps
+
+
Ichneumonidae
Gambrus ultimus (Cressson)
3
+
Mymaridae
Gonatocerus sp.
+
+
Eucoilidae
Trybliographa sp.
12
++
Kleidotoma sp.
+
+
Rhoptromeris sp.
+
+
Diapriidae
Trichopria sp.
+
+
Encyrtidae
Adelencyrtus odonaspidis
+
+
(Fullaway)
Scelionidae
Telenomus sp
+
+
Sphecidae
Crabro rufibasis (Banks)
+
+
Platygastridae (1 species)
+
+
Dryinidae
Gonatopus sp.
+
+
* Braconidae, and Pteromalidae were determined by V. Gupta;
Diapriidae and Scelionidae by P. Marsh; Pompilidae,
Sphecidae, Dryinidae, and Eucoilidae by A. Menke; Eulophidae
and Mymaridae by M. Shaufer; and Formicidae by D. Williams.


No. Horn Flies Emerged / Trial (Mean + SE)
94
Fig. 4-7. Predation rates by four species of Philonthus on
horn flies under simulated field conditions. Different
letters indicate significant differences at P = 0.05
level (100 g manure: 100 horn fly eggs/replicate). N =
10, 3, 10, 7 and 14 for control, hepaticus. P.
f lavolimbatus. P^ ventralis and P_;_ loncricornis.


Table 3-3. Scarabaeidae collected
central Florida. The occurre
enumerated: (+) rare, (++) co
53
in pastures in north
nee codes for species not
mmon, and (+++) abundant.
Trapped
Extracted
Scarabaeinae
Onthophagus gazella F.
971
+++
O. taurus (Schreber)
71
++
0. pennsylvanicus Harold
300
+++
O. hecate blatchleyi Brown
259
+++
O. striatulus floridanus
1
(Beauvois)
0. oklahomensis Brown
165
++
O. tuberculifrons Harold
286
+++
Copris minutus (Drury)
7
+
Canthon pilularius (L.)
42
++
Boreocanthon probus (Germar)
1
Phanaeus vindex MacLeay
31
+
P. igneus floridanus d'Olsoufieff
4
Cetoniinae
Euphoria sepulchralis (F.)
4
Hybosorinae
Hybosorus illigeri Reiche
4
Aphodiinae
Ataenius gracilis (Melsheimer)
9
A. picinus Harold
55
++
A. fattigi Cartwright
2
A. imbricatus (Melsheimer)
39
+
A. platensis Blanchard
1
Aphodius campestris Blatchley
15
+
A. lividus (Olivier)
1077
+++
A. parcus Horn
15
+
A. fimetarius (L.)
22
+
A. cuniculus Chevrolat
8
Geotrupinae
Geotrupes egeriei Germar
4
Bradycinetulus ferrugineus
(Beauvois)
2
Dynastinae
Strategus splendens (Beauvois)
1
* Scarabaeidae were determined by
M. Thomas,
P. Skelley, P.
Choate and J. McNamara.


92
only (sterilized by freezing), all died on the 4-6th day
before they developed into the second instar. This
indicates that Philonthus larvae are obligate predators and
are unable to survive by coprophagy.
Under simulated field conditions. Adults of P.
f lavolimbatus. P. hepaticus. P. longicornis. and P.
ventralis were tested for their predatory effects under
simulated field conditions. All four species significantly
reduced the mean numbers of horn flies emerged when the
results were compared with the control, no beetles (F(441) =
29.59, P < 0.01; Fig. 4-7). Duncan's multiple range test
showed that P^ loncricornis caused significantly higher
mortality of the horn flies than was caused by any of the
remaining three species (P < 0.05). It destroyed 80.4% of
the flies emerged compared to the control. It was found
also to prey on the fly pupae and newly emerged adult flies.
The next most effective species was P^ ventralis. which
destroyed 54.8% of the horn flies merged (P < 0.05). It
was found also to prey on horn fly pupae. Pj_ flavolimbatus
and Pi hepaticus significantly (P <: 0.05) reduced the mean
number of emerging flies by 40.6% and 32.9%, respectively,
compared with the control, but no significant difference (P
> 0.05) was found between these two species in the mean
number of horn flies that emerged. When two beetles were
used for each trial, a reduction of 95.8%, 91.9% and 70.3%
of horn fly emergence was obtained for Pi. lonaicornis. P.


84
after the pats were prepared. RIFA workers could also
infest the cone-trap covered pats through underground
tunnels. RIFA has a multiple-queen system and its nest
consists of multiple mounds and underground tunnels (Oi et
al. 1994). The fire ants built up small new colonies in the
heavily-infested pats before the horn fly finished its
immature development. Damaged adult horn flies were
observed in the vial screwed on the top of the cone traps
covering the pats in both treatments and I attribute this
damage to RIFA. The quantitative effect of RIFA on horn fly
populations will be reported in the next chapter. Another
reason for poor survivorship in the field was possibly that
the laboratory colony had been maintained for a long time
leading to genetic changes during the fly's domestication
(Bartlett 1984), and flies were no longer adapted to the
field environment. Desiccation plus high ambient
temperature during the day might be the primary abiotic
factors which decrease immature horn fly survivorship in the
dung.
Predation Rates of Predators on Horn Fly Immatures
[
Philonthus spp.
Members of the genus Philonthiis have been tested as
effective predators of horn flies (Thomas and Morgan 1972b;
Harris and Oliver 1979; Roth 1982; Fincher and Summerlin


82
Results and Discussion
Laboratory Horn Fly Survival
Percentage hatches of horn fly eggs were from 78 to 94%
(avg. 85.7 4.1%) when the eggs were distributed on paper
towel strips which were inserted into the artificial pat for
24 hrs. The percentage survival of the eggs to the adult
stage was from 22 to 41% [29.8 6,15 (SE)].
Fauna-Caused Field Horn Fly Mortalities
1992 July trial
Horn flies started to emerge on the 9th day after their
eggs were seeded under the artificial pat. Most of the
flies emerged on the 10th and 11th day. A few emerged on
the 12th and 13th day. An average of 2.9 0.95 (SE) horn
flies emerged from the pats covered by emergence traps to
exclude other arthropods from arriving in the dung (range 0-
8); an average of 0.7 0.3 (SE) flies emerged from the pats
exposed to other arthropods (range 0-3). An independent
Student's t-test showed that the mean number of horn flies
that emerged from uncovered pats was significantly smaller
than that emerged from covered pats (t(18) = 2.51, P < 0.05;
Fig. 4-3). Horn fly numbers were reduced by an average of
75.86% in the uncovered pats compared with the covered ones.


153
Frank, J.H. 1976.
in Florida.
Platvstethus spidulus Er. (Staphylinidae)
Coleopt. Bull. 30: 157-158.
Frank, J.H. 1982. The parasites of the Staphylinidae
(Coleptera). Univ. Florida Agrie. Exp. Station
Bulletin 824, 118 pp.
Frank, J.H. 1991. Staphylinidae (St
Brathinidae, Empelidae). pp. 3
Immature Insects. Vol. 2. Kend
Dubuque, Iowa.
aphylinoidea) (including
41-352. In Stehr (ed.):
all/Hunt Publ. Co.,
Fullaway, D.T. 1921.
18: 219-221.
Horn fly contr
ol.
Hawaii For. Agrie.
Glaser, R.W. 1924. Rearing flies fo
with biological notes. J. Econ
r
experimental purposes
Entomol. 17: 486-497.
Graham, O.H. and J.L. Hourrigan. 1977. Eradication programs
for the arthropod parasites of livestock. J. Med.
Entomol. 13: 629-658.
Granett, P. and E.J. Hansens. 1956.
control on milk production. J.
467 .
The effect of biting fly
Econ. Entomol. 49: 465-
Granett, P. and E.J. Hansens. 1957.
the effect of biting fly contr
cattle. J. Econ. Entomol. 50:
Further observations on
ol on milk production on
332-336.
Greer, N.I. and J.F. Butler. 1973. Comparison of horn fly
development in manure of five animal species. Florida
Entomol. 56: 197-199.
Greer, N.I. and J.F. Butler. 1975.
regulators on the bionomics an
fly, Haematobia irritans (L.).
University of Florida, Gainesv
Effect of insect growth
d control of the horn
Ph.D. Dissertation,
ille, 103 pp.
Hafez, M. 1939. Some ecological observation on the insect
fauna of dung. Bull. Soc. Fouad Ier Entomol. 23: 241-
387.
Hall
R.D. and K.E. Doisy. 1989. Wa
control of horn flies (Diptera
cattle. J. Econ. Entomol. 82:
lk-through trap for
: Muscidae) on pastured
530-534.
Halliday, R.B. and E. Holm. 1987. Mites of the family
Macrochelidae as predators of two species of dung
breeding pest flies. Entomophaga 32: 333-338.


23
ectoparasitoids of fly
imaszewski and Blume
predator of larvae (Roth 1982). Three exotic species, P.
lonqicornis (cosmopolitan), P. flavocinctus (from southeast
Asia) and P. minutus (from Australia), were tested to reduce
horn fly emergence by 61.1%, 68.1% and 99.2%, respectively,
under laboratory conditions (Fincher and Summerlin 1994).
Eight species of Aleochara are known to occur in cattle
dung in the USA (Fincher 1990) and the adults prey on the
eggs and larvae of cyclorrhaphous Diptera. But in the
immature stages they are solitary
pupae within the host puparium (K~.
1986) .
Hvdrophilidae. Sixteen species of hydrophilid beetles
have been reported from cattle dung in the continental USA
(Fincher 1990). Members of the genus Cercvon are abundant,
but the adults and larvae are dung-feeders (Sanders and
Dobson 1966; Hafez 1939; Merritt 1976; Thomas and Morgan
1972b). Sphaeridium scarabaeoides (L.), which is very
common in dung, has been considered as an important horn fly
predator in larval (Bourne and Hays 1968; Hammer 1941;
Macqueen and Beirne 1975; Mohr 1943; Poorbaugh et al. 1968;
Sanders and Dobson 1966; Thomas and Morgan 1972b) and adult
stages (Hammer 1941). The larva showed the highest predation
rate on horn fly larvae at 80F, and no predation below 40F
under laboratory conditions (Bourne and Hays 1968).
Histeridae Twenty-two species of histerid beetles are
associated with cattle dung in the continental USA (Fincher


15
pesticide-impregnated ear tags were in use, widespread
insecticide resistance occurred throughout the country
(Sheppard 1990). Subsequent development of horn fly
resistance to other insecticides, especially pyrethroid
compounds (Quisenberry and Strohbehn 1984; Sheppard 1984;
Sheppard and Joyce 1994; Kunz and Schmidt 1985), has
decreased field efficiency of chemical insecticides used for
horn fly control. In addition, insecticides used for feed
through for control of dung-inhabiting flies leave a residue
in cattle dung, which adversely affects abundance of
biocontrol agents (e.g. Scarabaeidae and Staphylinidae) of
the horn fly in pastures (Fincher 1992; Madsen et al. 1990;
Wall and Strong 1987).
Genetic. Physical and Immunological Control
Sterile male technique has been used for horn fly
control (Eschle et al. 1977; Kunz and Eschle 1971) and was
successful. The difficulties for keeping this technique
continue to work are isolation of the target area (Kunz et
al. 1974), raising large numbers of competitive sterile
flies, and difficulty in combining several control measures
(Graham and Hourrigan 1977).
A practical horn fly walk-through trap designed by
Bruce (1938) has been tested to control the horn fly. This
trap controlled an average of 54-73% and 50% horn flies on
cattle in Missouri (Hall and Doisy 1989) and Texas (Bruce


CHAPTER 1
INTRODUCTION
The horn fly, Haematobia irritans (L.), is an Old World
species which invaded the United States before 1886. Its
populations grew and spread, and reached Florida by 1891.
Its range is from Canada to Brazil (Hargett and Goulding
1962b; Moya-Borja 1990) and it can be a serious problem for
the cattle industry in the New World wherever it occurs.
Adult male and female flies are hematophagous, and bite
frequently (Harris et al. 1974). Cattle are their primary
hosts and fly numbers on a single animal can reach thousands
(Bruce 1964; Butler 1975, 1990; Fay 1986; Kinzer 1970). The
blood loss and annoyance can cause substantial reduction of
milk production and live weight gain in domestic cattle
(Bruce and Decker 1947; Harvey and Brethour 1979).
The fly larvae are coprophagous and develop only in
fresh cattle dung (Hogsette and Koehler 1986; Macqueen and
Beirne 1975; Skidmore 1991). Studies have shown that other
insects present in cattle dung are important mortality
factors for the immature stages of the horn fly (Blume et
al. 1970; Fay 1986; Fay and Doube 1983; Fay et al. 1990;
Harris 1981; Harris and Blume 1986; Kunz et al. 1972;
Legner 1986; Macqueen and Beirne
1
1975; Sanders and Dobson


colony maintained by Dr. J. F. But!
115
er (chapter 4). The eggs
used in the test were collected fro
m egg trays within 4
hours after deposition, suspended i
n well water, and
pipetted on paper towel strips (6 c
m long X 2 cm wide). The
paper strips then were placed under
the edge of the
artificial pads. Artificial pats
were positioned every 15
m in a grid pattern in the Amdro-tr
eated and the control
area, none closer than 20 m from th
e edge.
The mean number (avg SD) of
RIFA and horn flies
collected from the Amdro treated ar
ea and the control area
were compared by an independent Stu
dent's t-test (Sigmaplot
1994). Mean numbers (avg SD) of
selected epigean or dung-
inhabiting arthropod taxa were comp
ared by the general
linear model procedure (GLM; SAS In
stitute 1990). The
numbers of arthropods per trap or p
at were transformed by
log (n + 1) to reduce heteroscedast
icity (Dowdy and Wearden
1983) before analysis of Student's
t-test or GLM.
Results
RIFA Population Control
In 1992, Amdro was applied on
October 1 when fire ant
populations were not significantly
different between the
Amdro-treated and control area. Te
n days after Amdro was
applied, averages of 123.3 118.4
and 663.2 310.3


Table 3-5 (continued)
Trapped
Extracted
Ceratopogonidae
Forcipomyia brevipennis (Macquart;
) ++
63
Scatopsidae (1 species)
+
42
Tachinidae (2 species)
+
13
Psychodidae (2 species)
+
129
Sciaridae
Bradysia coprophili (Lintner)
++
107
Phoridae
Megaselia sp.
+
16
Anisopodidae
Sylvicola notialis Stone
++
266
Chloropidae
Monochaetoscinella nigriconls (Loew) ++
32
Drosophilidae (1 species)
++
44
Tabanidae (1 species)
+
4
Tipulidae (1 species)
5
4
* Muscidae were determined by D. Deonier, G. Gagne, and G.
Steck; Sarcophagidae by G. Steck and G. Dahlem; Bibionidae,
Sepsidae and Sphaeroceridae by G. Steck; Cecidomyiidae and
Sciaridae by J. Gagn and W. Grogan; Phoridae by D. Deonier;
Ceratopogonidae by W. Grogan and G. Steck; Dolichopodidae by
K. Ahlmark; Empididae and Chloropidae by G. Steck; and
Anisopodidae and Ephydridae by W. Wirth.


83
horn flies emerged from
1992 August trial
An average of 3.0 0.56 (SE)
the pats covered by emergence traps to exclude other
arthropods from arriving in the dung (range 0-5); an
average of 1.0 0.26 (SE) flies emerged from the pats
exposed to other arthropods (range
Student's t-test showed that the mean number of horn flies
emerged from uncovered pats was significantly smaller than
that emerged from covered pats (t(18)
0-2). An independent
3.25, P < 0.01; Fig.
4-3). Horn fly numbers were reduced by 66.7% in the
uncovered pats compared with those in the covered pats.
Combined data from July and August 1992 showed that the dung
arthropod community caused 71.26% mortality to immature horn
flies in artificially formed cowpats in north-central
Florida.
The mortality of immature horn flies caused by the dung
fauna in the present study was lower than mortalities given
in previous reports. Contributions of the arthropod
community in pastures to horn fly mortality were reported as
87.9% (Roth 1989) and 90% (Kunz et al. 1972) in Texas, 97.7%
in Missouri (Thomas and Morgan 1972b), and 79-84% in
Australia (Fay et al. 1990).
Horn fly survivorship was lower in the field (covered
pats) than in the laboratory. One reason was infestation by
the red imported fire ant (RIFA), which arrived within 24


50
efficient predators of the horn fly (Roth 1982; Harris and
Oliver 1979; Fincher and Summerlin 1994). Except for P.
hepaticus, local species (R ventralis, P. sericans and P.
rectanqulus) were less commonly collected than the two
adventive ones. This indicates that these adventive species
are well established and competed successfully with native
Philonthus species.
Aleochara notula was very abundant during the survey.
The adults prey on fly eggs and larvae of cyclorrhaphous
Diptera, and its immature stages are solitary
ectoparasitoids of fly pupae within the host puparia
(Klimaszewski and Blume 1986; Harris and Blume 1986). They
were collected from dung in emergence boxes from the time
the dung was placed in the boxes until about 3 weeks
thereafter. A sarcophagid [Raydnia delicta (Walker)] was
found to be the main host of this beetle in the dung,
followed by Brontaea debilis (Williston) and B^ cilfera
(Malloch) [Muscidae]. No horn fly pupae were found to be
parasitoidized by Aleochara notula. Twenty-three A^_ notula
were reared from Ravinia delicta pupae in one cowpat in
August 1993. The adults of A_¡_ notula were found to be
abundant and active in fresh to one week old dung.
Oxvtelus incisus and Atheta sp. were abundant in the
survey. They may play an important role in reducing horn
fly populations in north central Florida if they prey on
horn fly immatures. Valiela (1974) cited Oxvtelus


1969; Thomas and Morgan 1972b). The horn fly viability is
affected by predation, parasitoidism, and competition for
the same food source (Harris and Blume 1986). Important
horn fly predators are reported to be Staphylinidae,
Hydrophilidae, Histeridae (Harris and Blume 1986; Thomas and
Morgan 1972b), and the red imported fire ant (Howard and
Oliver 1978; Schmidt 1984; Summerlin et al. 1984b). The most
important competitors are dung beetles (Scarabaeidae)
(Fincher 1986; Anderson and Loomis 1978; Blume et al. 1973).
The most abundant parasitoids are Spalanoia spp. (Harris and
Blume 1986).
Although the parasitoids of horn flies have been
investigated in Florida (Butler et al. 1981; Escher 1977),
the arthropod community in cattle dung has not been examined
quantitatively, and little attention has ever been paid to
its species composition. Such information, however, is very
important for evaluation of the biotic mortality of horn fly
immature stages, and for determination of the need for
introduction of biocontrol agents for horn fly control.
The primary goal of the present study was to provide
extensive information on the species composition and
abundance of horn fly natural enemies and their effect on
the horn fly immatures in north central Florida. In detail,
the objectives of this study were 1) to determine the
composition and abundance of the arthropod community in
cattle dung; (2) to monitor seasonal distribution of these


108
1982, 1990) under simulated field
present study the adults of ft coen
conditions. In the
osus were found to prey
primarily on horn fly eggs. Each
of 27.1 20.78 fly eggs (range 12
2.3 2.05 fly larvae (range 0-7)
beetles penetrated into and stayed
Petri dishes, and came out to feed
larvae of this species also were ob
horn fly eggs and larvae too.
Carabidae.
Aspidoqlossa subanqulata adults were observed to prey
on horn fly eggs. Remnants of fly larvae were not observed
in the Petri dishes. This beetle plrobably consumed all the
beetle consumed an average
53) and an average of
per day (n = 15). These
in the cotton ball in the
during the night. The
served to prey on the
fly larvae. Only chewed small piec
left in Petri dishes. Stenocrepis g
es of egg chorions were
uatuordecimstriata was
observed to consume 150-200 horn fl
battlefield-like situation was left
eggs chewed by these beetles could
The results show that the medium an
beetles inhabiting dung will kill f
whenever they encounter them. They
y eggs a day. Because a
in the Petri dishes,
not be counted exactly,
d small-sized carabid
ly eggs and larvae
are able to consume most
of the eggs and all the larval tissue.


34
for food or space as rare among coprophagous Dptera.
In Australia, horn flies developing in cattle dung
experienced little competition for food or losses due to
predation from other arthropods (Bornemissza 1960, 1970).
The effect of other coprophagous flies on the horn fly
population in North American pastures has not been reported.
Parasitoids
The Hymenoptera in dung, with the exception of ants,
are parasitoids. Blume (1985) listed 43 species of
parasitoid Hymenoptera associated with cattle dung. Twenty-
two of them, summarized by Fincher (1990), have been
reported as parasitoids of the horn fly in the United States
(Lindquist 1936; Thomas 1981; Thomas and Morgan 1972a; Figg
et al. 1983; Watts and Combs 1977; Combs and Hoelscher 1969;
Harris and Summerlin 1984; Schreiber and Campbell 1986).
Depner (1968) and Peck (1974) reported nine parasitoid
species from Canada. Although many species of parasitoids
have been reported to attack the horn fly, the most common
parasites are in the genus Spalangia (Harris and Blume
1986).
Some staphylinid beetles have been recorded to
parasitoidize the pupae of horn flies. They are Aleochara
bimaculata Gravenhorst (Klimaszewski 1984), and Aleochara
sp. (Cervenka and Moon 1991). In these species the newly
hatched larva enters a fly puparium, develops within, and


144
Horn Fly Control
Horn fly control has been based on chemicals,
especially the advent of pesticide-pregnated ear tags
(Byford et al. 1985; Harvey and Brethour 1970, 1979;
Quisenberry and Strohbehn 1984; Schmidt and Kunz 1980), but
chemicals cause resistance of the fly (Quisenberry and
Strohbehn 1984; Sheppard 1984, 1990; Sheppard and Joyce
1994; Kunz and Schmidt 1985), pollution to the environment,
and reduction of natural enemies in the dung (by feed
through) (Fincher 1992; Madsen et al. 1990; Wall and Strong
1987). Moreover, chemical control is costly and of limited
value because of the continuous availability of dung as
habitat (Legner 1986) .
The horn fly is an immigrant pest in North America and
Australia and plagued their cattle industry. However, it
rarely reaches pest status in its originEurope and Africa.
Therefore classical biological con
trol may be the best
strategy for control of the horn fly.
To import more scarabs to the pastures may not be the
best strategy. Release and establishment of imported dung-
burying scarab beetles has not reduced pest fly densities in
pastures in Hawaii, Texas, California and Florida in the
U.S., and Australia. There is even
negative effect of populations of
predators by interrupting their habitat (Legner 1986), and
some indication of a
imported scarabs on native


36
and in establishment of programs in selected areas of the
country.
Hunter et al. (1991) reported 22 species of
Staphylinidae in their survey in Texas and described the
pattern of seasonal distribution and diel flight activity of
abundant species. Hanski and Hammond (1986) observed that
predatory species were more patchily distributed than
saprophagous species. Successes in rearing Staphylinidae
have been reported in North America (Mank 1923; Harris and
Oliver 1979; Hunter et al. 1986b) and in Europe (Hinton
1944; Paulian 1941) and Africa (Tajwfik et al. 1976a, b, c) .
Life history and habits have
been observed of some
species of Philonthus (Mank 1923;
Hunter et al. 1986a;
Tawfik 1976a,b,c), Platystethus (H
inton 1944; Legner and
Moore 1977) and Aleochara (Peschke
and Fuldner 1977; White
and Legner 1966; Drea 1966). Dung
-inhabiting Staphylinidae
have four stages: egg, larva, pupa
and adult. There are
three larval instars for most subf
amilies and two for
Paederinae (Frank 1991). The eggs
are white, some with
sculptures in the surface of chori
on, and hatch within 2-5
days. An egg burster was observed
for the embryo within the
egg (Hinton 1944). The neonates of
Aleochara gnaw a hole in
a fly pupa, enter and develop in t
le puparia (Peschke and
Fuldner 1977; White and Legner 196
5; Drea 1966), and the
remaining genera are free living a
nd pupate in the dung or
soil under the dung pats (Hinton 1
944; Mank 1923; Frank


49
number of individuals, followed by Scarabaeidae. The numbers
of Hydrophilidae, Carabidae and Histeridae were much lower.
Staphvlinidae. An estimated 10,000 adult Staphylinidae
were collected and over 7000 were identified to species
level. The specimens belong to seven subfamilies, of which
Staphylininae, Aleocharinae, Oxytelinae and Paederinae were
most abundant (Table 3-2), Xantholininae were common, and
Tachyporinae and Osorinae were rare. Oxvtelus incisus and
Aleochara notula were abundant from trapping and extracting,
Tinotus spp. were abundant in extractions, and Acrotona
hebeticornis and Atheta sp. were abundant in pitfall traps.
Anotvlus spp., Philonthus flavolimbatus. P. lonqicornis and
Neohypnus pusillus were common from trapping or extracting.
Platvstethus americanus. Philonthus hepaticus were common in
extraction. Platvstethus spiculus were only collected by
extracting and Platvdracus tomentosus were only collected by
trapping.
Members of the genus Philonthus have been well
documented to be predators of fly immature stages (Hammer
1941; Harris and Oliver 1979; Hunter et al. 1989; Macqueen
and Beirne 1975; Roth 1982; Thomas and Morgan 1972b). Six
Philonthus species were collected in this study. P.
hepaticus. P. flavolimbatus. and P_¡_ lonqicornis were more
common than the other three species, of which P^ rectanqulus
was rare. The adventive species P. lonqicornis and P.
flavolimbatus were common and have been reported to be


CHAPTER 2
LITERATURE REVIEW
History and Importance of the Horn Fly
History
The horn fly, Haeraatobia irritans (L. ) is a
widespread, economically important pest of cattle (Morgan
and Thomas 1974, 1977). It hitchhiked into the United States
from southern France before 1886, but before then was known
only from Europe (Riley 1889). This fly is believed to have
entered the United States on cattle through the port of
Philadelphia and was first collected in Camden, New Jersey,
in August 1887 (Bruce 1964; Morgan and Thomas 1974). It
spread rapidly in North America (Hargett and Goulding 1962b;
Stone et al. 1965; Butler 1990), reaching Florida by 1891
(Osborne 1896). It reached South America later, and is
still moving south (Moya-Borja 1990). This fly has occupied
almost the whole of Brazil, Paraguay, Uruguay and Bolivia,
and over half of Argentina (Foil and Hogsette 1994).
Economic Importance
Horn flies are conspicuous because they can occur on
the host in large numbers. It has been estimated that
10,000 flies may occur on a single animal (Bruce 1964).
4


112
1984b). However, there have been no data to show the impact
of RIFA on horn flies and other arthropods in pastures of
Florida.
During the survey of the arthropod community inhabiting
cattle dung in north central Florida, high populations of S.
invicta were found to infest cowpat
colonies were observed to develop i
after the pats were deposited, and
observed to carry horn fly larvae t
objective of the present study was
of RIFA on populations of horn flie
a RIFA-infested pasture versus a pa
in north central Florida.
s in pastures. New
n cowpats within a week
RIFA workers were
o their nests. The
to determine the impact
s and other arthropods in
sture treated with Amdro
Materials and Methods
Field tests were conducted dur
1993 in a pasture south of Gainesvi
Florida. There were 40 cattle in t
experiment was conducted. Two plot
were delineated: one was used as th
other as the control. The plots we
separating them, but cattle had acc
gates were left open. No horn fly
during the year 1992 and 1993 by th
RIFA populations were controll
ing the fall of 1992 and
lie, Alachua Co.,
he pasture when the
s (about 2 acre each)
e treatment field and the
re adjacent with a fence
ess to both plots because
control was implemented
e herd owner,
ed by using a fire ant


75
procedure of Thomas and Morgan (1972b). Horn fly eggs less
than 4 hours old were removed from the egg collection tray,
suspended in well water and pipetted on paper towel strips
moistened with well water. Twenty-five eggs were counted on
each towel strip. A metal hoop, 20.32 cm diameter and 5.08
cm high, was placed on a section of grass in a large metal
pan. The area within the hoop was moistened with well
water. One-hundred eggs [25 eggs on each of four paper
strips (6 cm long X 2 cm wide)] were placed on the grass
within the hoop, each strip being placed at each of four
side of the edge. The area within the hoop was covered with
manure, the hoop was removed and a simulated manure pat was
formed (approximately 25 cm diam X 5 cm high). The
simulated manure pats were held in a rearing room for 7-8
days and then covered with cone traps to collect emerged
horn flies. Six replicates were conducted.
Field studies on horn fly mortality
Field mortality of the horn fly was evaluated during
July and August 1992 at pasture A where an arthropod survey
was conducted at the same time (chapter 3). Horn fly eggs
from the laboratory colony were used. Manure collection and
artificial pat formation were performed according to the
procedure for laboratory survival studies.
Two trials were conducted, one each in July and August.
Twenty simulated manure pats for each trial were formed


124
area (Table 5-3) in 1992. Eh_ flavo
effective predator of the horn fly
1994; Roth 1982; Harris and Oliver
Amdro-treated area compared with th
Anotylus insicrnitus and A_^_ nanus we
Amdro-treated area and are also pre
tested in chapter 4 for A¡. insigni
Other staphylinids including A
Acrotona hebeticornis were common i
area compared with the control area
Aleochara notula is a predator of h
the adult stage and a parasitoid of
immature stage (Klimaszewski and B1
Acrotona hebeticornis on flies is n
Pitfall trapping of both years
were the most common hydrophilid sp
fimetarius the most common scarabae
sp. the most common carabid species
limbatus. also an
(Fincher and Summerlin
1979), is common in the
e control area in 1993.
re also common in the
dators of the horn fly as
tus.
leochara notula. and
n the Amdro-treatment
in 1993 (Table 5-5).
orn eggs and larvae as
fly pupae as the
ume 1986). The effect of
ot yet known.
showed that Cercvon spp.
ecies; CK. gazella and A_;_
id species; and Tachvs
RIFA Effect on Dung-Inhabiting Arthropods
1992 trial
Staphylinids and hydrophilids were commonly extracted
from dung pats bought into the laboratory, but the mean
numbers of these two families of Coleptera were not
significantly different (P > 0.05, Table 5-6).


12
clustering of flies on horns is no longer as common as in
previous times. (Foil and Hogsette 1994). The flies
congregate mainly on the shoulders and sides of the animal
(Hogsette and Koehler 1986), because these places are least
disturbed by the swishing of the tail, tossing of the head
and kicking of the legs (Harvey and Launchbaugh 1982). Horn
flies alter their distribution (from back and shoulder area
to belly area) on cattle treated with pyrethroid-impregnated
ear tags in comparison with untreated cattle (Byford et al.
1987) .
In a series of experiments, Hargett and Goulding
(1962b) found that many horn flies, dislodged during the
night, failed to find the host until daylight. They
concluded that upon emerging, horn flies depend more on
vision than on olfactory or heat stimuli to find a host. In
contrast, Kinzer et al. (1978), using an artificial cow as a
field attractant, found that temperature and C02 were prime
factors in horn fly orientation. Dalton et al. (1978) found
that radiated heat rather than internal temperature was
involved and that cow odor was the most powerful influence
on responding flies, especially at close range.
Color and breed are reported to affect the horn fly's
host preference. Horn flies prefer dark-colored animals
(Bruce 1964). Brahman cattle were less attractive than
European breeds (Tugwell et al. 1969). Calves and yearlings
with long, thick hair are not as desirable as are cattle


Parasitoids 34
Ecology and Biology of Staphylinidae 35
3. The ARTHROPOD COMMUNITY IN NORTH CENTRAL
FLORIDA PASTURES 38
Materials and Methods 39
Results and Discussion 48
Coleptera 48
Staphylinidae 49
Scarabaeidae 55
Carabidae 57
Hydrophilidae 60
Histeridae 60
Diptera 61
Hymenoptera 6 5
Other Insects and Animals 67
4. EFFECT OF ARTHROPOD PREDATORS ON HORN FLY
SURVIVORSHIP IN PASTURES AND UNDER LABORATORY
CONDITIONS 73
Materials and Methods 74
Test of the Arthropod Community on
Horn Fly Survivorship 74
Laboratory studies on horn fly
survivorship 74
Field studies on horn fly
mortality 75
Determination of Predation Rates of
Predators 77
Collection and colonization of
predators 77
Predation rate test under
laboratory conditions ... 77
Predation rate test under
simulated field conditions 78
Results and Discussion 82
Laboratory Horn Fly Survival .... 82
Fauna Caused Horn Fly Mortalities 82
1992 July trial 82
1992 August trial 83
Predation Rates of Predators on Horn
Fly Immatures 84
Philonthus spp 84
Under laboratory conditions 86
Under simulated field
conditions 92
Oxytelinae 96
Paederinae 100
Xantholininae 100
Tachyporinae 105
v


52
collected sporadically.
Diel activity was also different among the species. A.
notula and Philonthus spp. were mainly collected during the
day time, but Tinotus spp., Acrotona hebeticornis. Atheta
sp. and Oxvtelus incisus were mainly collected during the
night.
Adult Staphylinidae began to leave the dung placed in
the emergence boxes a few hours after the dung was brought
into the laboratory. The number increased the next day, was
most abundant within about a week, and then declined. Most
of the staphylinid beetles left the boxes within two weeks
after dung was collected and placed in them. Those
extracted later developed from the immature stages present
in the dung when it was collected.
Successful biological control of dung-inhabiting pest
flies depends on ecological factors, especially seasonal
distribution and the habitat preference of natural enemies.
In north central Florida, horn flies are active all year
round, without overwintering (Wilkerson 1974), and become
most active from May to October. Staphylinidae are the most
important predators of the horn fly (Fincher 1990; Harris
and Blume 1986) elsewhere, but our survey results showed
that the numbers of Staphylinidae did not reach their peak
until July, especially Philonthus spp. and A^_ notula (Fig.
3-4) which have biocontrol potential for the horn fly.
Because of this 'lag' time, numbers of horn flies increase


LIST OF TABLES
Table Page
3-1. Summary of invertebrates collected in pastures in
north-central Florida from 1991 to 1993 42
3-2. Staphylinidae collected in pastures in north central
Florida 45
3-3. Scarabaeidae collected in pastures in north central
Florida. 53
3-4. Coleptera collected in pastures in north central
Florida, with exclusion of Staphylinidae and
Scarabaeidae 58
3-5. Diptera collected in pastures of north central
Florida 62
3-6. Hymenoptera collected in pastures in north central
Florida 66
3-7. Miscellaneous invertebrates collected in pastures in
north central Florida 68
4-1. Larval predation rates of five Philonthus species
during the whole developmental period on horn fly
eggs and larvae under laboratory conditions .... 91
4-2. Predation by N. pusillus larvae and adults on eggs
and larvae of horn flies under laboratory
conditions 102
5-1. Numbers of horn flies emerged from artificial pats
in the Amdro-treated and control areas, October
1992 and 1993 119
5-2. Numbers of arthropod specimens collected by pitfall
traps from the Amdro-treated and control areas in
October 1992 120
vii


7
conducted, the breed and sex of animals, and the size of the
horn fly population (Steelman 1976).
Horn flies serve as vectors for a filarial nematode,
Stephanofilaria stilesi. and a bacterial disease, mastitis,
between cattle. The nematode reduces the value of hides
for leather and causes blemishes that are a problem when
registered animals are used for exhibition purposes
(Steelman 1976). Mastitis may render dairy cattle useless.
Horn flies are also potential vectors of other diseases
(Butler 1975) .
Biology of the Horn fly
Life History
The horn fly belongs to the Animal Kingdom Arthropoda
-Atelocerata (Uniramia) Hexapoda (Insecta) Pterygota -
Diptera Aristocera Muscoidea r Muscidae Haematobia
irritans (Linnaeus) (Borror et al. 1989). It is a
holometabolous insect, with four stages: egg, larva, pupa
and adult. After the host has been found, the adult horn
fly leaves the vicinity of the host only to oviposit on
fresh manure. Eggs hatch within 24 hours and the larvae
pass through three instars, pupating in the third instar
skin in 3 to 5 days. It takes nine days from oviposition to
adult emerge at 30C (Melvin and Beck 1931). The field
developmental time depends on the temperature (Hargett and
Goulding 1962a; Lysyk 1992a&b; Wilkerson 1974).


122
than that from the control area (1.
08 1.17). Horn fly
numbers were reduced by 62.9% from
the pats in the control
area compared with those from the p
ats in the Amdro-treated
area.
RIFA Effect on Epicjeal Arthropods
1992 trial
Specimens of staphylinids were
abundantly collected in
the traps 4 weeks after Amdro was a
pplied. The mean number
of these insects collected in the P
mdro-treated area was
significantly greater than that in
the control area (F(1 18) =
5.24, P < 0.05; Table 5-2). Mean n
umbers of hydrophilids,
carabids, and scarabs were low in t
oth the Amdro-treated and
control area, but the mean number o
f carabids collected in
the Amdro-treated area were signifi
cantly greater than that
in the control area (P < 0.05; Tabl
e 5-2). On the contrary,
the mean number of scarabs collecte
d in the control area was
significantly greater than that in
the Amdro-treated area
(F(1 18) = 3.49, P < 0.05) Mean num
bers of mites and
collembolans collected in both area
s were not significantly
different (P > 0.05).
1993 trial
Because it took more time to c
pntrol the fire ant
populations in the pasture by apply
ing the bait Amdro,


the original mounds are disturbed
1983). All the mounds in the Amdrci
with flags and the area around each
with Amdro. After another two week
developed small mounds were found a
thereafter, horn fly eggs were seed,
in the field and the epigean arthrc
RIFA Effect on the Horn F1Y
When fire ant workers were confined in Petri dishes and
exposed to horn fly larvae, they attacked these fly larvae
immediately (Fig. 5-1). From the dung pats exposed 8 days
117
(Williams and Lofgren
-applied area were marked
mound was re-treated
s when only a few newly
nd treated with Amdro
ed in the artificial pats
pods were collected.
in October 1992, an average of 1.94
emerged per pat in the Amdro-treate
only 0.11 0.31 flies emerged per
(Table 5-1). An independent Studen
the mean number of horn flies that
Amdro-treated area was significantl
emerged from pats in the control ar
Table 5-1). Horn fly numbers were
pats in the control area compared w
in the Amdro-treated area. The fie
1993 showed that the number of horn
emerged from the artificially seede
treated area was significantly grea
2.17 horn flies
d area; an average of
pat in the control area
t's t-test showed that
emerged from pats in the
y greater than that
ea (t= 3.44, P < 0.01;
reduced by 94.3% from the
ith those from the pats
Id test data in October
flies (2.91 0.57)
d pats in the Amdro-
ter (t = 2.63, P < 0.05)


coverall average was 71.3%.
Predation rates were tested under laboratory (Petri
dishes) and simulated field conditions (test cages
containing artificially formed pats) with adults of five
field-collected Philonthus species against horn fly
immatures. IL, longicornis had the highest predation rate.
Aleochara notula was very abundant during the survey and its
adult is an effective predator of horn fly immatures.
Fifteen other staphylinid, one hydrophilid, one histerid,
and two carabid species were also found to prey on horn fly
eggs and larvae. Larvae of eight staphylinid species and
one tenebrionid species were also found to prey on horn fly
immatures.
Red imported fire ants (RIFA) were observed to infest
cowpats heavily all year round and prey on horn fly larvae
and pupae in the pastures. RIFA caused 94.3% and 62.9%
mortality of horn flies, respectively, for October 1992 and
1993, compared with a RIFA-controlled area in the field,
though it reduced populations of horn fly natural enemies.
xii


BIOGRAPHICAL SKETCH
Guanye Hu is a native of China and was born on May 15,
1954. He started primary education at Dahu Elementary
School in 1960. After he worked as a tractor driver for two
years, he went to Taishan Medical College in 1974 and
studied medicine. The author started his career at the same
college as an assistant lecturer teaching medical
parasitology and entomology in 1978. He spent three years
on his M.S. in parasitology and entomology at Shanxi Medical
College between 1982-1984. Then he returned to Taishan
Medical College and was appointed as an assistant professor
and associate chair of the Department of Parasitology and
Entomology. He came to the USA as
a visiting scientist
working at the Florida Medical Entcmology Laboratory on
mosquito ecology in 1989 and began the doctoral program at
the University of Florida in the spring term of 1991. He is
married to Yanfen Chen, and they have a daughter, Wenli, and
a son, Jesse.
170


Table 3-1. Summary of invertebrates collected by two methods
in pastures
in north-central
Florida from 1991-1993.
Class and Order
No. Families
No. Species
Insecta
Orthoptera
2
3
Homoptera
3
5
Coleptera
14
109
Diptera
20
35
Hemiptera
5
9
Lepidoptera
3
3
Hymenoptera
16
24
Dermaptera
1
1
Thysanoptera
1
1
Isoptera
1
1
Collembola
3
15
Arachnida
Acari
5
12
Araneae
1
4
Diplopoda
1
1
Nematomorpha
1
1
Nematoda
1
1
Mollusca
1
1
Total
79
226


101
and larvae for food, without manure, predation on eggs and
larvae occurred. One consumed 8 eggs and 6 larvae per day;
the other consumed 2 eggs and 9 larvae per day.
Lithocharodes ruficollis. The adult ruficollis
preyed on both horn fly eggs and larvae. Each adult beetle
(n = 5) consumed an average of 2.Q 2.1 (SD) eggs (range
0-6) and an average of 3.2 0.4 (SD) larvae (range 3-4)
per day.
Neohypnus. Adults of three species of Neohvpnus were
collected during the survey. In the present study, adults
of all three species were observed to prey on horn fly eggs
and larvae. Each N. attenuatus consumed an average of 4.8
1.3 fly eggs (range 3-6) and an average of 4.0 1.58
larvae (range 2-6) per beetle per day (n = 4). Each N.
emmesus consumed an average of 5.4 1.74 horn fly eggs
(range 3-8) and an average of 4.0 1.1 fly larvae (range
3-6) per beetle per day (n = 5).
Each adult N. pusillus consumed 4.2 eggs and 5.9 larvae
of the horn fly daily under laboratory conditions (Table 4-
2). The adult beetle was observed to hold the anterior end
of a fly larva or the side of a fly egg in its mouth and
then move quickly to get under the cotton ball. Five
females of rt pusillus were observed to lay eggs in the
Petri dishes and two of them oviposited daily for 2-3 weeks
when they were provided horn fly eggs and larvae for food,
without manure. The newly hatched larvae preyed on horn fly


103
O
CO
~o
0
Tj
0
E
3
W
c
o
O
>*
0
Q_
5 7 9 11 13 15
Predator Larval Age (Day)
Fig. 4-9. Daily predation rates of N. pusillus larvae
(n = 19) on horn fly eggs and larvae under laboratory
conditions.


20
the first 24 hours are very important for reducing the horn
fly populations (Fay et al. 1990; Blume et al. 1970).
Effect of the Whole Insect Fauna on Horn Fly Production
The presence of other arthropods in cattle dung has
been shown to reduce populations of horn flies (Blume et
al. 1970; Fay 1986; Fay and Doube 1983; Harris and Blume
1986; Kunz et al. 1970, 1972; Macqueen and Beirne 1975; Roth
1989; Sanders and Dobson 1969; Thomas and Morgan 1972b).
Blume et al. (1970) showed a significant negative
correlation between the mean numbers of horn flies and the
mean numbers of insects of other species produced per dung
pat. Roth (1989) reported that dung fauna caused an average
of 87.9% mortality in immature horn flies in naturally
infected dung pats between April to mid-September in Texas.
In a similar study in Missouri, Thomas and Morgan (1972b)
reported 97.7% mortality of horn flies in exposed dung pats.
When dung pats were covered within five minutes to eliminate
competition from other insects, an average of 66.5 flies
emerged from each pat (Kunz et al. 1970); but when pats were
not covered and competition from other insects was allowed,
an average of 6.6 flies emerged per pat (Kunz et al. 1972).
In Florida, Wilkerson (1974) and Greer and Butler (1973)
found an average of 8 to 19 flies emerging from each manure
pat in the summer months.
In Australia, Fay et al. (1990) reported that fauna-


Aleocharinae 105
Hydrophilidae 107
Histeridae 107
Carabidae 108
Tenebrionidae 109
Anthicidae 109
5. EFFECT OF THE RED IMPORTED FIRE ANT (RIFA) ON THE
HORN FLY AND OTHER ARTHROPODS IN PASTURES . Ill
Materials and Methods 112
Results 115
RIFA Population Control 115
RIFA Effect on the Horn Fly .... 117
RIFA Effect on Epigeal Arthropods 122
1992 trial 122
1993 trial 122
RIFA Effect on Dung-Inhabiting
Arthropods 124
1992 trial 124
1993 trial 127
Discussion 134
6. CONCLUSION AND GENERAL DISCUSSION 137
REFERENCE LIST 14 6
BIOGRAPHICAL SKETCH 170
vi


55
rapidly in the spring before populations of Staphylinidae
increase accordingly. If predator abundance could be
increased and maintained at a high level during this period
by inundative releases of native or introduced species with
similar activity patterns, horn fly numbers might be
reduced.
Scarabaeidae. Over 8,000 scarab beetles were collected
and more than 5,000 were identified to species (Table 3-3).
They belong to 27 species of six subfamilies, of which
Scarabaeinae (12 species) and Aphodiinae (10 species) were
predominant. Onthophacrus qazella.
were the most abundant (Table 3-3), followed by 0.
pennsvlvanicus. 0. h. blatchlevi. 0. oklahomensis and O.
tuberculifrons. O. taurus, Canthon pilularius. Phanaeus
and Aphodius lividus
vindex. and Ataenius picinus were commonly collected.
Unlike Staphylinidae, very few Scarabaeidae were
collected from January to March. Their numbers started to
increase in April, the first peak was in May, and the second
and third in July and November 1993, respectively (Fig. 3-
2) The peak in May was mainly contributed by A^_ lividus;
the one in July by CK. qazella and supplemented by A.
lividusand the November peak by pennsvlvanicus. O. h.
blatchlevi. O. oklahomensis. and 0^_ tuberculifrons (Fig. 3-
5)
High numbers of A^_ lividus in the early summer (May
1993) contributed to the breaking of the dung pats. These


157
Kinzer, H.G., J.W. Reeves and J.W.
location by the horn fly: Fiel
artificial device for measurin
stimuli. Environ. Entomol. 7:
Atmar. 1978.
d attraction
g attraction
375-378.
Host
of an
to various
Klimaszewski, J. 1984. A revision
Gravenhorst of America north
Staphylinidae, Aleocharinae)
No. 129.
of the genus Aleochara
of Mexico (Coleptera:
Mem. Entomol. Soc. Can.
Klimaszewski, J. and R.R. Blume. 19
Aleochara verna Say and Aleoch
(Coleptera: Staphylinidae, A1
Bull. 40: 32.
86. New host records for
ara notula Erichson
eocharinae). Coleopt.
Knipling, E.F and W.C.
livestock, p. 167.
McDuffie. 1956. Flies that affect
USDA. Year Book Agrie. 591 pp.
Koehler, P.G. and J.F. Butler. 1976
Protection Pointers. IFAS. 18,
Horn flies.
10 pp.
Livestock
Koehler, P.G. and J.F. Butler. 1980. Control of the external
parasites with forced-use dust bags. Livestock
Protection Pointers.
Koehler, P.G. and J.F. Butler. 1984. Horn fly resistance to
synthetic pyrethroid ear tags. Livestock Protection
Pointers, No.7, 14 pp.
Koskela, H. 1972. Habitat selection
Staphylinidae (Coleptera) in
dung. Ann. Zoo. Fennici 9: 156
of dung-inhabiting
relation to age of the
-171.
Krafsur, E.S. and C.M. Ernst. 1983.
compositions and reproductive
populations, Haematobia irrita
Muscidae) in Iowa, USA. J. Med
Physiological age
biology of horn fly
ns irritans (Diptera:
Entomol. 20: 664-669
Krantz, G.W. 1983. Mites as biologi
dung-breeding flies, with spec
Macrochelidae. pp. 91-98. In
and L. Knutson (eds.): Biologi
Mites. University California,
cal control agents of
ial reference to the
M.A. Hoy, G.L. Cunningham
cal Control of Pests by
Berkeley.
Krombein, K.V., P.D. Hurd and D.R.
Hymenoptera in America north o
Institute, Washington, D.C. Vo
Smith. 1979. Catalog of
f Mexico, Smithsonian
1. 1, 2, pp. 1-2209.
Kunz, S.E. 1978. Notes on the seasonal activity of dung
infesting Diptera in central Texas. Southwest. Entomol.
3: 167-170.


CHAPTER 4
EFFECT OF ARTHROPOD PREDATORS ON HORN FLY SURVIVORSHIP
IN PASTURES AND UNDER LABORATORY CONDITIONS
The results of an arthropod community survey in north
central Florida (chapter 3) showed that over 220 species of
invertebrates are present in association with dung. The
dung arthropod community has been shown to reduce horn fly
populations in the USA (Blume et al. 1970; Thomas and Morgan
1972b; Kunz et al. 1972; Roth 1989), Canada (Macqueen and
Beirne 1975) and Australia (Fay et al. 1986, 1990). Insect
predators are reported as the primary biotic mortality
factor of the horn fly (Thomas and Morgan 1972b; Macgueen
and Beirne 1975; Harris and Blume 1986; Roth 1983, 1989).
Studies have shown that the most important predators are
coleopterans of the families Staphylinidae, Hydrophilidae,
and Histeridae (Bornemissza 1968; Fay 1986; Legner 1986;
Roth 1989; Sanders and Dobson 1969; Thomas and Morgan
1972b). Philonthus beetles (Staphylinidae) have been shown
to have high potential for predation on horn fly immatures
(Roth 1982; Fincher and Summerlin 1994; Harris and Oliver
1979). In Florida, only Escher (1977) and Butler et al.
(1981) reported parasitism of horn
been no studies of the effect of the whole arthropod
73
flies, but there have


77
Determination of Predation Rates of Predators
Collection and colonization of predators
Adults of predatory beetles were extracted using the
emergence cages (Fig. 3-1) from the pats collected in
pastures A and B. Colonies of predatory beetles were
initiated and maintained by following the method of Hunter
et al. (1986b) for Philonthus. Five to 10 pairs of beetles
were placed in 5 cm high X 15.24 cm diameter plastic
containers with several moist paper towels crumpled one on
top of the other. Two circular holes (2.5 cm diam) were cut
through the lid; one was covered by fine cloth screen to
allow for a free flow of air; the other was open for
insertion of predators and prey (horn fly eggs). Horn fly
eggs (500-1000) suspended in distilled water were pipetted
on the surface of the towels twice a week as a food source.
Predation rate test under laboratory conditions
For observation of predation, predators were confined
individually in Petri dishes (5.5 cm diameter X 1.3 cm
high), lined on the bottom with a moist paper towel. A
water-soaked cotton ball was provided for humidity, and
horn fly eggs and 1st instar larvae were provided for food.
An observation cage as described by Hinton (1944) was also
made for rearing P. americanus in the laboratory. The cage
consisted of a well cut in a piece of styrofoam. The well


120
Table 5-2. Numbers of arthropod specimens collected by
pitfall traps from the Amdro-treated and control area
in October 1992 (n = 10).
Arthropod taxa Amdro
Control
Staphylinidae
Mean
5.0
1.4
SD
4.51
1.11
Range
0-16
0-3
Significance
F = 5.
24 P
<
0.05
Hydrophilidae
Mean
0.2
0.3
SD
0.4
0.46
Range
0-2
0-3
Significance
F = 0.
24, P
>
0.10
Scarabaeidae
Mean
1.0
3.4
SD
1.0
3.78
Range
0-10
0-34
Significance
F = 3 .
49 P
<
0.10
Carabidae
Mean
0.2
0
SD
0.4
0
Range
0-2
0
Significance
F = 2.
25, P
<
0.10
Mites
Mean
12.9
4.5
SD
26.82
6.56
Range
0-88
0-23
Significance
F = 0,
P >
0.
10
Collembola
Mean
19.5
120.4
SD
21.9
234.7
Range
0-64
0-800
Significance
F = 1.
25, P
>
0.10


147
Blume, R.R. 1970. Insects associated with bovine droppings
in Kerr and Bexar Counties, Texas. J. Econ. Entomol.
63: 1023-1024.
Blume, R.R. 1984. Euoniticellus
Scarabaeidae): description
biology of adults. Environ.
int
of
En
ermedius (Coleptera:
adults and immatures and
tomol. 13: 1064-1068.
Blume, R.R. 1985. A checklist, distributional records, and
annotated bibliography of the
bovine droppings on pastures in America north of
Mexico. Southwest. Entomol.
insects associated with
1-55 .
Blume, R.R. and A. Aga. 1978. Onthdphagus qazella: Progress
of experimental releases in south Texas. Folia Entomol.
Mex. 39-40: 190-191.
Blume, R.R., J.M. Matter and J.L. E
qazella: Effect on survival of
laboratory. Environ. Entomol.
schle. 1973. Onthoohagus
horn flies in the
2: 811-813.
Blume R.R., S.E. Kunz, B.F. Hogan a
Biological and ecological inve
in central Texas: Influence of
manure. J. Econ. Entomol. 63:
nd J.J. Matter. 1970.
stigations of horn flies
other insects in cattle
1121-1123.
Bolton, H.T. 1980. The role of sem
behavior of the horn fly, Haem
(Diptera: Muscidae). Diss. Dep
Univ. Florida. 212 pp.
iochemicals in the
atobia irritans (L.),
t. Entomol. & Nematol
/
Bornemissza, G.F. 1960. Could dung
our pastures? J. Aust. Inst.
eating insects improve
Agrie. Sci. 26: 54-56.
Bornemissza, G.F. 1968. Studies on
Pachylister chinensis in Fiji
the control of buffalo-fly in
16: 673-688.
the histerid beetle
and its possible value in
Australia. Aust. J. Zool.
Bornemissza, G.F. 1970. Insectary
dung breeding flies by the act
Onthophaqus qazella F.(Coleopt
Aust. Entomol. Soc. 9: 31-41.
studies on the control of
ivity of the dung beetle,
era: Scarabaeinae) J.
Bornemissza, G.
1965-1975.
F. 1976.
Aust.
The Australian dung beetle project
Meat Res. Committee Rev. 30: 1-30.
Borror, D.J., C.A. Triplehorn and N
Introduction to the Study of I
Saunders Company, Pheladelphia
.F. Johnson. 1989. An
nsects (6th ed.).
, PA, 875 pp.


130
Table 5-6. Continued.
Arthropod taxa
Sciaridae
Anisopodidae
Ceratopogonidae
Psychodidae
Hymenoptera
(except RIFA)
No. collected
Amdro
Mean
SD
Range
11.3
28.74
0-126
F = 0.02,
Control
17.3
34.21
0-104
P > 0.05
Mean
SD
Range
0.06
0.23
0-1
F = 0.69
0.2
0.71
0-3
P > 0.05
Mean
SD
Range
1.9
4.35
0-17
0.2
0.71
0-4
P > 0.05
Mean
SD
Range
2.8
4.35
0-15
F =
9.77,
0
0
0
P < 0.01
Mean
SD
Range
0.9
2.3
0-10
F =
1.15,
0.4
1.11
0-4
P > 0.05


156
Howard, F.W. 1975. Arthropod popula
treated with Mirex baited to s
ant populations. Ph.D. Diss. L
Baton Rouge.
tion dynamics in pastures
uppress red imported fire
ouisiana State Univ.,
Howard, F.W. and A.D. Oliver. 1978.
permanent pastures treated and
red imported fire ant control.
901-903.
Arthropod populations in
untreated with Mirex for
Environ. Entomol. 7:
Hunter, J.S. Ill, D.E.
of Staphylinidae
Burleson County,
Bay and G.T.
associated wi
Texas. Southw
Fincher. 1986a. A survey
th cattle droppings in
est. Entomol. 11: 83-88.
Hunter, J.S. Ill, D.E. Bay and G.T.
Laboratory and field observati
and habits of Philonthus cruen
flavolimbatus.
Fincher. 1989.
ons on the life history
tatus and Philonthus
Southwest. Entomol. 14: 41-47.
Hunter, J.S. Ill, D.E. Bay, G.T. Fi
Beerwinkle. 1991. Seasonal dis
activity of Staphylinidae (Col
wooded pasture in east central
ncher and K.R.
tribution and diel flight
eoptera) in open and
Texas. J. Kans. Entomol.
64: 163-173.
Hunter, J.S. Ill and G.T. Fincher.
records for the Afro-Asian dun
qazella (Coleptera: Scarabaei
20: 24-25.
1985. Five new state
g beetle Onthoohagus
dae). J. Entomol. Sci.
Hunter, J.S. Ill, G.T. Fincher and
for rearing Philonthus spp. as
droppings. J. Entomol. Sci. 21
D.E. Bay. 1986b. Methods
sociated with cattle
: 83-86.
Jones, D.J. and W.L. Sterling. 1979. Manipulation of red
imported fire ants in a trap crop for boll weevil.
Environ. Entomol. 8: 1073-1077.
Kenlin, R.L. and P.G. Alingham. 199
reponse of cattle exposed to
irritans exigua). Vet. Parasitol. 43: 115-129.
2. Acquired immune
uffalo fly (Haematobia
Kinzer, H.G. 1970. Ground applicati
Malathion and Fenthion for hor
Mexico. J. Econ. Entomol. 63:
on of ultra-low-volume
n fly control in New
736-739.
Kinzer, H.G. and J.M. Reeves. 1974.
location of the horn fly. Envi
Dispersal and host
ron. Entomol. 3: 107-111.


133
Table 5-8. Numbers of staphylinid s
Amdro-treatment and control ar
pecimens extracted from
ea in October 1993.
Subfamily
Species Name
Amdro
Control
Oxytelinae
Oxytelus incisus
995
338
Apocellus sphaericol
lis 3
3
Paederinae
Rugilus angularis
2
0
Lithocharis sororcul
a 10
4
Thinocharis sp.
10
6
Staphylininae
Philonthus hepaticus
47
29
P. flavolimbatus
22
0
P. longicornis
6
2
P. ventralis
19
4
Xantholininae
Neohypnus pusillus
21
5
Aleocharinae
Acrotona hebeticorni
S 17
12
Atheta sp.
25
18
Tinotus amplus
138
9
T. brunnipes
62
97
Aleochara notula
1
0
Others
27
31
Total
1361
558


11
only on fresh cattle droppings (Butler 1975; Kunz et al.
1970; Sanders and Dobson 1969; Skidmore 1991). Droppings
older than 10 minutes are unattractive unless the crust is
broken (Bruce 1964). The flies move from the shoulders of
cattle to the area near the tail, as animals deposit manure
(Foil and Hogsette 1994). Possibly the gravid female
receives a signal of impending defecation with the lifting
of the tail of the host, and is ready to oviposit on the
fresh manure as soon as it hits the ground, or even before
the feces hit the ground (Skidmore 1991). Mohr (1943)
observed that all adult horn flies left the dung pats two
minutes after the dung was deposited. Each female lays an
average of 24 eggs per batch and 15 batches during its
lifetime (Bruce 1964). The eggs are laid singly or in
groups of 4 to 6 (Hogsette and Koehler 1986), and a total of
200 to 400 is laid during the life time of the female
(Wilkerson 1974).
Host Orientation and Location
Adult flies remain on the body of cattle day and night,
unless the females leave for oviposition (Hammer 1941;
Hargett and Goulding 1962b; Kunz et al. 1970; Morgan
1964), though they make short rapid flight from one part of
the host to another, or between adjacent hosts (Bruce 1964).
The name 'horn fly' originates from the tendency of these
flies to cluster at the base of horns of the host, but


141
insects including
central Florida; and 5) the 'lag' time, because horn flies
are R-selected insects, with higher fecundity and
developmental rate, but beneficial
predators, competitors and parasites, are k-selected
insects, with lower fecundity and developmental rate.
Effect of Red Imported Fire Ants on Horn Fly Survival
Red imported fire ants (RIFA) were observed to infest
cattle pats heavily all year round and to prey on horn fly
larvae and pupae in the pastures. RIFA caused 94.3% and
62.9% mortality of immature horn flies in cattle pats during
the fall of 1992 and 1993, respectively, compared with
cattle pats from which RIFA were excluded. RIFA has also
been reported to reduce other pest
fields (Adams et al. 1981; Reagan et
Sterling 1979) and in pastures (Bruc
Pimentel 1955; Summerlin and Kunz 1
intensive control of RIFA may indire
populations, such as horn flies.
populations in crop
t al. 1972; Jones and
ce 1964; Laurence 1954;
978). Therefore,
ectly liberate pest
RIFA has adverse effect on oth
er beneficial insects
associated with the horn fly in cattle dung. This is
because RIFA is a general predator. It preys whatever they
meet. They may prey on the immature predators (e.g.
staphylinids) or competitors (e.g. scarabaeids and flies).
However, the effect of RIFA on horn flies are


121
Table 5-3. Numbers of staphylinid specimens collected by
pitfall traps from the Amdro--t
in October 1993 (n = 10).
.reated and
control area
Subfamily
Species Name
Amdro
Control
Oxytelinae
Anotylus nanus
10
0
A. insignitus
1
0
Paederinae
Rugilus angularis
1
0
Staphylininae
Philonthus hepaticus
1
0
P. flavolimbatus
7
1
P. longicornis
23
4
Xantholininae
Neohypnus attenuatus
0
1
Aleocharina
Acrotona hebeticorni
s 1
5
Atheta sp.
5
0
Tinotus brunnipes
1
1
Aleochara notula
0
1
Tachyporinae
Bryoporus rufescens
0
1
Total
50
14


1994). During the field survey of
86
the arthropod community
in the present study, specimens of
six species of Philonthus
were found in the dung. Living specimens of five species
were colonized in the laboratory for predation rate tests:
P. flavolimbatus, P. hepaticus, P.
lonaicornis, P. sericans
and P. ventralis.
Under laboratorv conditions.
Adults of all five
Philonthus species preved on horn :
:ly eggs and early instar
larvae. P. lonaicornis (Fia. 4-4)
preferred fly eggs to
larvae; P. ventralis preferred lar^
rae to eggs; the other
three species preyed about equally
on eggs and larvae (Fig.
4-5). P. lonaicornis was observed
to consume as many as 6-7
horn fly eggs per minute without ii
iterruption; it chewed
the fly egg chorion into shapeless
pieces while ingesting
the fluid. Victimized fly larvae 1
iad only the
cephaloskeleton and the sclerotizec
1 structures of the
posterior spiracle (peritreme, button and spiracular slits)
left on the paper towel or on the
;otton ball in the Petri
dish. ANOVA showed that predation
rates by these five
Philonthus species on combinations
of horn fly eggs and
larvae were significantly difieren)
' (4, 136) = 259.2 P <
0.01; Fig. 4-5). The predation ra)
;e of P. lonaicornis
(120.1 /day) was significantly higl
er (P < 0.05) than those
of the remaining four species, followed bv P. ventralis
(38.9/day). The predation rates ol
: P. flavolimbatus, P.
sericans and P. hepaticus were not
significantly different


44
J FMAMJ JASOND
Month 1993
Fig. 3-3. Seasonal changes in number of species of
Staphylinidae and Scarabaeidae in pastures of north
central Florida.


164
Schmidt, C.D. 1984. Influence of fi
re ants on horn flies and
other dung-breeding Diptera in
Bexar County, Texas.
Southwest. Entomol. 9: 174-177

Schmidt, C.D. and S.E. Kunz. 1980.
Fenvalerate and stirofos
ear tags for control of horn f
lies in range cattle.
Southwest. Entomol. 5: 202-206

Schmidt, C.D., C.R. Ward and J.L. E
schle. 1972. Rearing and
biology of horn fly in the lab
oratory: Effect of
density on survival and fecund
Entomol. 2: 223-224.
ity of adults. Environ.
Schoenly, K. 1983. Arthropods assoc
iated with bovine and
equine dung in an ungrazed chi
huahuan desert ecosystem.
Ann. Entomol. Soc. Am. 76: 790
-796.
Schreiber, E.T. and J.B. Campbell.
1986. Parasites of the
horn fly in western Nebraska.
211-215.
Southwest. Entomol. 11:
Schreiber, E.T., J.B. Campbell, D.J
. Boxler and J.J.
Petersen. 1987a. Comparison of
beetles collected from
the dung of cattle untreated a
nd treated with
fenvalerate ear tags and pastu
red on two range types in
western Nebraska. Environ. En
tomol. 16: 1135-1140.
Schreiber, E.T., J.B. Campbell, S.E
. Kunz, D.C. Clanton and
D.B. Hudson. 1987b. Effect of
horn fly (Diptera:
Muscidae) control on cows and
gastrointestinal worm
(Nematoda: Trichostrongylidae)
treatment for calves on
cow and calf weight gains. J.
454.
Econ. Entomol. 80: 451-
Sheppard, D.C. 1984. Fenvalerate an
d flucythrinate
resistance in horn fly populat
1: 305-310.
ions. J. Agrie. Entomol.
Sheppard, D.C. 1990. Insecticide re
sistance in horn flies in
USA. pp. 1216-1222. In 16th Wo
Symposium. Brazil. 1263 pp.
rid Buiatrics Congress
Sheppard, D.C. 1994. Dispersal of w
ild-captured, marked horn
flies (Diptera: Muscidae). Env
34 .
iron. Entomol. 23: 29
Sheppard, D.C. and J.A. Joyce. 1994
. High levels of
pyrethroid resistance in horn
flies (Diptera: Muscidae)
selected with cyhalothrin. J.
1593 .
Econ. Entomol. 85: 1587-


13
with short, thin hair (Hammer 1941). Bulls attract and hold
more horn flies than cows, because of the effect of male sex
hormone, testosterone (Dobson et al. 1970). Though the horn
fly's principal host is cattle, it sometimes attacks other
animals such as goats, horses, mules, deer, dogs and,
rarely, man, especially when the bovine host is absent
(Bruce 1964).
Diapause and Dispersal
Hargett and Goulding (1962b) reported that the horn fly
overwinters in the southern United States as an adult, but
in the northern United States and Canada it overwinters as a
diapausing third instar larva or as a pupa. There has been
no observed diapause in horn flies in Florida (Wilkerson
1974). Horn fly diapause is mediated in the fall by
decrease of temperature and photoperiod (Thomas et al. 1987)
and is terminated in the spring by increasing temperatures
of the substrates (Lysyk 1992b; Thomas et al. 1987).
Some authors stated that horn fly adults are sedentary
on the host, but leave for oviposition on manure pats (Bruce
1942, 1964; McLintock and Depner 1954). On the contrary,
Chamberlain (1981, 1982) and Hoelscher et al. (1968) found
that horn flies moved several hundred meters from a host.
Moreover, Kinzer and Reeves (1974) and Tugwell et al. (1966)
showed the potential for long distance dispersal of natural
horn fly populations. Horn flies were found to be able to


Table 5-6. Numbers
of specimens
of
arthropod
taxa
collected
per cowpat sample from the
Amc
Iro-treated area and
control area,
October 1992
(n
= 18) .
Arthropod taxa
No. collected
Amdro
Control
Staphylinidae
Mean
8.7
23.4
SD
10.46
53.77
Range
0-32
0-236
F = 0.28,
P
>
0.05
Hydrophilidae
Mean
2.9
1.8
SD
4.0
3.37
Range
0-17
0-10
F = 2.01,
P
>
0.05
Muscidae (except H_
. irritans)
Mean
3.9
0.6
SD
4.74
1.8
Range
0-16
0-7
F = 8.5,
P <
0
. 01
Sarcophagidae
Mean
3.6
1.6
SD
5.21
4.23
Range
0-21
0-14
F = 4.53,
P
<
0.05
Sepsidae
Mean
6.4
1.7
SD
21.11
5.18
Range
0-93
0-22
F = 1.17,
P
>
0.05
Sphaeroceridae
Mean
29.4
14.2
SD
42.7
16.92
Range
0-170
0-47
F = 1.32,
P
>
0.05
Cecidomyiidae
Mean
5.7
9.1
SD
8.25
11.26
Range
0-30
0-46
F = 1.19,
P
>
0.05
129


No. Scarabaeidae Collected / Trap
54
24
8
0
3
Aphodius lividus
Ataenius imbricatus
At. picinus
nciijLd jriTl rji
XL
0
24
Phanaeus vindex
Canthon pilularius
I

12 i
8
4
0
Onthophagus gazella
O. h. blatchleyi
O. pennsylvanicus
0. oklahomensis
O. tuberculifrons
O. taurus
T*
F
T"
M
rrrj.iTTi
Ji
K
/
M J J A
Month 1993
S O N D
Fig. 3-5. Seasonal distribution of scarabaeid species in
pasture A, collected by trapping. N = 10 X 2 for May to
October and N = 10 for the remaining months.


165
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Smith, J.P., R.D. Hall and G.D. Tho
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Steelman, C.D. 1976. Effects of ext
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Stewart, T.B. and D.R. Davis. 1967
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Stone, A., C.W. Sabrosky, W.W. Wirt
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Summerlin, J.W., D.E. Bay, R.L. Har
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Summerlin, J.W., D.E. Bay, R.L. Har
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No. Horn Fly Immatures Consumed / Predator Larva
90
Larval Development (day)
Fig. 4-6. Daily predation rates of 5 species Philonthus
larvae on horn fly eggs and larvae under laboratory
conditions. N = 33, 22, 35, 6 and 29 for P.
lonqicornis. P. flavolimbatus. P. sericans, P.
hepaticus and P^ ventralis. respectively.
40
30
20
10
0
40
30
20
10
0


149
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Byford, R.L., S.S. Quisenberry, T.C
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Haematobia irritans (Diptera:
Entorno1. 78: 768-773.
Campbell, J.B. 1976. Effect of horn
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Econ. Entomol. 69: 711-712.
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Integrated pest
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orally resistant horn
attle treated with
s. Environ. Entomol. 16:
Sparks and J.A.
secticide cross
tant populations of
Muscidae). J. Econ.
fly control on cows as
g weights on calves. J.
Cervenka,V.J. and R.D.
with fresh cattle
Entomol. Soc. 64:
Moon. 1991.
dung in Min
131-145.
of the fly problems, pp.
Scuda (eds.). Rural
? ARD/IANR, Univ.
11. 317 pp.
Arthropods associated
nesota. J. Kansas
Chamberlain, W.F. 1981. Dispersal
host proximity. Southwest. Ent
Chamberlain, W.F. 1982. Dispersal
Wildflies. Southwest. Entomol
Cheng, T.H. 1958. The effect of bit
gain in beef cattle. J. Econ.
Collins, H., A. M. Callcott, T.C. L
1992. Seasonal trends in effe
hydramethylnon (Amdro) and fen
control of red imported fire a
Formicidae). J. Econ. Entomol.
of horn flies: Effect of
omol. 6: 316-325.
of horn flies: II.
7: 230-234.
ing fly control on weight
Entomol. 51: 275-278.
ockley and A. Ladner,
ctiveness of
oxycarb (LOGIC) for
nts (Hymenoptera:
85: 2131-2137.
Combs, R.L. and C.E. Hoelscher. 196
parasitoids found associated w
northeast Mississippi. J. Econ
9. Hymenopterous pupal
ith the horn fly in
Entomol. 62: 1234-1235.
Cutkomp, L.K and A.L. Harvey. 1958. The weight responses of
beef cattle in relation to control of horn and stable
flies. J. Econ. Entomol. 51: 72-75.


125
Table 5-4. Numbers of arthropod specimens collected by
pitfall traps from the Amdro-t.reated and control area
in October 1993 (n = 10).
Arthropod taxa
Amdro
Control
Staphylinidae
Mean
60.4
16.8
SD
27.88
10.82
Range
9-106
2-44
Significance
F = 15.86, P <
0.01
Hydrophi1idae
Mean
2.2
0.4
SD
1.89
0.49
Range
0-5
0-1
Significance
F = 6.71, P <
0.05
Scarabaeidae
Mean
2.0
2.2
SD
2.6
2.14
Range
0-8
0-6
Significance
F = 0.25, P >
0.10
Carabidae
Mean
0.3
0.1
SD
0.64
0.3
Range
0-2
0-1
Significance
F = 0.60, P >
0.10
Fall armyworm
Mean
2.9
0.1
SD
2.17
0.3
Range
0-6
0-1
Significance
F = 15.79, P <
0.01


106
8) per day. Each T. amplus (n = 1C
>) consumed an average of
1.69 0.85 (SD) fly larvae (range
1-4) per dav. Atheta sp.
females laid eggs when they were pr
-ovided horn fly larvae
and eggs for food without manure.
Aleochara notula has been
reported as a predator on horn fly
immatures (Klimaszewski
and Blume 1986; Harris and Blume IS
*86). In the present
study each adult A. notula Cn = 29;
Fig. 4-10) was observed
to consume an average of 40.3 IS
.83 horn fly eggs (range
10-82) and an average of 4.7 2.
8 larvae (range 2-12) per
day. The results showed that adult
:s of this species preyed
primarily on horn fly eggs. The fe
pales of A. notula laid
eggs daily when they were provided
horn fly eggs and larvae
for food, without manure. When A_.
notula adults were
tested under simulated field condit
:ions (100 g manure and
200 horn fly eggs/one beetle) it claused a 32% reduction of
horn fly emergence compared to cont
rols. The population of
A. notula was verv hiah in the fiel
d (chapter 3), so this
beetle may be a very important pred
ator of horn flies in
north central Florida.
Larvae of Atheta were observed
to prey on horn fly
larvae. Each larva (n=7) preyed on
2.36 0.64 horn fly
larvae per day and on 20.3 9.68 f
ly larvae during the
whole development period (11-13 day
s). All the larvae died
without pupating, indicating that they might need cow dung
to make chambers for pupation as did EL. americanus. A.
insicrnitus and 0. incisus.


109
Tenebrionidae
Gondwanocripticus obsoletus ad
ults were commonly
collected, but were not observed to
prey on horn fly eggs
and larvae. All the adults died af
ter a few days when they
were confined in Petri dishes and p
rovided horn fly eggs and
larvae, without manure. One of its
larvae extracted alive
from the dung was confined in a Pet
ri dish, and provided
horn fly eggs and larvae for food,
without manure. It
consumed an average of 5.0 3.16
horn fly eggs (range 1-
25) and 1.4 0.49 larvae (range 1J
7) per day. It fed
actively for 5 days and became tran
quil for a day before
pupation.
Anthicidae
Anthicus sp. was not observed
to prey on horn fly eggs
and larvae. When five adults of th
is beetle were confined
in the Petri dishes and offered hor
n fly eggs and larvae for
food, without manure, they died in
a few days without
consuming any horn fly immatures.
Adults of two species of carab
id and larvae of one
species of tenebrionid have not pre
viously been reported as
predators of horn fly immatures, so
they are newly reported
here.
Present studies show that the
predatory beetles in dung
are diverse. Many of these species
were commonly collected


25
from bovine feces in a
that damages crops and attacks livestock, poultry and
wildlife (Hays and Hays 1959; Lyle and Fortune 1948). RIFA,
however, has been reported to prey on dung-inhabiting
immature Diptera (Laurence 1954; Bruce 1964). It invades
fresh dung pats which are less than 10 minutes old
(Summerlin et al. 1984a) and causes significant reduction of
horn flies in field studies (Howard 1975; Howard and Oliver
1978; Summerlin et al. 1984b; Schmidt 1984). Howard and
Oliver (1978) and Schmidt (1984) observed that fire ants
carried larvae and pupae of the horn fly from the pats.
Summerlin et al. (1977) showed that RIFA reduced emergence
of adult horn flies about 20 fold
laboratory study in Alabama; Schmidt (1984) showed a seven
fold increase of the horn fly in a pasture in Texas where
RIFA was controlled. Howard and Oliver (1978) found that 2-
5 times more horn fly pupae were recovered from pasture in
Louisiana where RIFA was controlled by Mirex bait than in
pasture where it was not controlled. Lemke and Kissam
(1988) reported that the number of horn flies emerging from
manure piles where RIFA was controlled using Pro-Drone was
55% greater than the number that emerged from piles in a
RIFA-infested field in South Carolina.
On the other hand, some evidence shows that RIFA not
only affects pest fly populationsJ but also has negative
effects on some other beneficial insects inhabiting the
dung, such as the scarab beetle Onthophagus gazella F.


46
Table 3-2. (continued)
Trapped Extracted
Aleocharinae
Acrotona hebeticornis Notman 336
Atheta sp. 207
Aleochara notula Erichson 507
Gnypeta floridana Casey ? 2
Tinotus amplus Notman 59
T. brunnipes Notman 76
Thyasophila sp. 2
Falagria dissecta Erichson 2
Meronera venustula (Erichson) 3
Hoplandria sp. 1 1
sp. 2 1
Thecturota sp. 1
Alaobia scapularis (Sahlberg) 1
Athetini (genus?) 6
Tachyporinae
Bryoporus rufescens LeConte 4
Mycetoporus flavicollis 1
(LeConte)
59
93
+++
179
191
+
+
+
Undetermined 265 5
* all Staphylinidae were determined by J. H. Frank.


CHAPTER 3
THE ARTHROPOD COMMUNITY IN NORTH CENTRAL FLORIDA PASTURES
The presence of other insects in cattle dung has been
shown to reduce populations of the horn fly, Haematobia
irritans (L.) (Blume et al. 1970, Thomas and Morgan 1972b,
Macqueen and Beirne 1975). Hence, a basic study of the
insect fauna is essential for evaluation of regulation of
the horn fly and for consideration of further need of
introduction of biocontrol agents into an area. Several
comprehensive surveys have been conducted in North America
(Macqueen and Beirne 1975; Merritt and Sanders 1977; Mohr
1943; Sanders and Dobson 1966; Poorbaugh et al. 1968; Blume
1970; Valiela 1969b; Wingo et al. 1974), and over 400
species of arthropods have been collected in or on cattle
dung in the United States (Harris and Blume 1986). No
comprehensive study of the fauna of cattle dung has yet been
reported for Florida. Arthropod species that affect horn
flies vary geographically, therefore, biotic mortality of
horn fly immature stages in Florida may not be assessed
using the data of arthropod composition and abundance from
any other states. Results reported herein are a study on
the arthropod community in pastures in north central
Florida. The objectives of this study were to determine the
38


161
Morgan, C.E. and G.D. Thomas. 1974.
of the horn fly, Haematobia ir
references on the buffalo fly,
and other species belonging to
U.S. Dept. Agrie. Mise. Pub. 1
Annotated bibliography
ritans (L.), including
H. exigua (de Meijere),
the genus Haematobia.
278, 134 pp.
Morgan, C.E. and G.D. Thomas. 1977.
bibliography of the horn fly,
including reference in the buf
Meijere), and other species be
Haematobia. U.S. Dept. Agrie.
Supplement I: Annotated
Haematobia irritans (L.),
falo fly, It exigua (de
longing to the genus
Mise. Publ. 1278, 38 pp.
Morgan, N.O. 1964. Autecology of th
Haematobia irritans (L.). Ecol
e adult horn fly,
ogy 45: 728-736.
Morgan, N.O. and O.H. Graham. 1966. Influence of cattle
diet on survival of horn fly larvae. J. Econ. Entomol.
59, 835-837.
Morgan, N.O. and C.D. Schmidt. 1966
eggs of the horn fly. J. Econ.
Variations in color of
Entomol. 59: 882-884.
Moya-Borja, G.E. 1990. A mosca do c
Distribuico ecologia e mtodo
combate. XVI Congresso Mundial
Congresso Latino Americano de
hifre na America Latina:
s alternativos de
de Buiatria, VI
Buiatria. pp. 1206-1208.
Oi,
D.H., R.M. Pereira, J.L. Stimac
Field applications of Beauveri
the red imported fire ant (Hym
Econ. Entomol. 87: 623-630.
and L.A. Wood. 1994.
a bassiana for control of
enoptera: Formicidae). J.
Okine, J.S. 1991. Aspects of oogene
Haematobia irritans (Linnaeus)
Thesis. Univ. Florida, Gainsvi
sis in the horn fly,
(Diptera: Muscidae).
lie, 135 pp.
Okine, J.S. and J.F. Butler. 1995.
plasma and erythrocyte diets o
Haematobia irritans (L.) (Dipt
and oogenesis. J.Agrie. Entorno
Effect of bovine blood
n adult horn fly
era: Muscidae) mortality
1. (in press).
Osborne, H. 1896. Insects affecting
Div. Entomol. Bull. 5: 114-121
domestic
animals.
USDA
Palmer, W.A. and D.E. Bay. 1981. A review of the economic
importance of the horn fly, Haematobia irritans (L.).
Protection Entomol. 3: 237-244.
Palomino, P., F.L. and W.E. Dale L.
spiculus (Coleptera: Staphyli
de mosca domestica. Revista Pe
1989. Platvstethus
nidae) predator de huevos
ruana Entomol. 31: 39-45.


142
significant (chapter 5). Millions of dollars has been used
to control RIFA as a pest in Florida. The actual economic
loss caused by RIFA is only $ 1 million. Without RIFA, the
horn fly populations would have been much higher than their
present status.
Effect of Predatory Beetles on Horn Flv Immatures
Predation rates by predators o
n immature horn flies
were tested under laboratory (Petri
dishes) and simulated
field conditions (test cages containing artificially formed
pats). The species shown to be pre
dacious on immature horn
flies include five Philonthus soeci
es. Aleochara notula. and
fifteen other staphylinid species;
Sohaeridium lunatum
(Hvdrophilidae); Hister coenosus (H
isteridae); and
Aspidocrlossa subancrulata and Stenoc
repis auatuordecimstriata
(Carabidae). Larvae of 5 staphylin
id species and one
tenebrionid species were also found
to prey on immature horn
flies. Among these tested species,
two Philonthus species.
S. lunatum, and H. coenosus have be
en reported previously as
predators of the horn fly, but all
the remaining species are
for the first time shown here to be
predators of immature
horn flies.
In the past, introduction of s
oarab beetles have been
considered to be a very important m
ethod for horn fly
control. Present study shows that
predators have greater


57
brought into the laboratory, most Scarabaeidae left the dung
placed in the emergence boxes within two days.
The results showed that the numbers of scarab beetles
built up later in the year than the numbers of horn flies.
Introduction of more efficient coleopteran competitors or
inundative releases of native species in the spring may
reduce the horn fly numbers in north central Florida.
Carabidae. A total of 20 species of carabids was
collected, but none were abundant. However, Tachvs sp.,
Ardistomis viridis, Stenocrepis quatuordecimstriata and
Tetragonoderus intersectus were common (Table 3-4). Some
large-sized species did not occur in the dung at all, such
as Calosoma sayi and Pasimachus spp., even though Pasimachus
spp. have been reported to be associated with bovine
droppings (Blume 1985). Some small- and medium-sized
species were commonly extracted from the dung. Since they
are general predators, they may be important in reducing
horn fly populations. Mohr (1943) and Hammer (1941)
reported, respectively that larvae of Carabidae were
predators of fly larvae in dung. Only two Galerita larvae
were collected in the dung, but mainy adult Carabidae
extracted developed from immature stages in the dung.
Most species in this family were collected
sporadically. The number of individuals was low all the
year and there were no obvious peaks (Fig. 3-2). Tachvs
sp., and S_j_ quatuordecimstriata were collected in most


CHAPTER 5
EFFECT OF THE RED IMPORTED FIRE
ANT ON THE HORN FLY
AND OTHER ARTHROPODS
IN PASTURES
The red imported fire ant (RII
A), SolenoDsis invicta
Burn (Hymenoptera: Formicidae), ir
vaded the United States
50 year ago from South America and
has spread across the
southern U.S. from Texas to North C
arolina (Lemke and Kissam
1988; Porter and Savignano 1990).
Though it is condemned as
a pest species for damaging some ac
rricultural crops,
livestock, and wildlife (Adams et e
1. 1976; Hays and Hays
1959; Lyle and Fortune 1948; Smitt
.le et al. 1983), RIFA has
been reported to be an important pr
edator of sugarcane
borers (Adams et al. 1981; Reagan e
it al. 1972), boll weevils
and bollworms (Jones and Sterling 1
979), and dung-inhabiting
immature flies (Bruce 1964; Laurenc
:e 1954; Pimentel 1955;
Summerlin and Kunz 1978).
RIFA invades fresh dung pats v
hich are less than 10
minutes old (Summerlin et al. 1984c
), and the workers were
observed to carry horn fly larvae e
nd pupae from the pats
(Howard and Oliver 1978; Schmidt 19
84). Studies elsewhere
show that the ants cause significar
t reduction of horn flies
in field studies (Bruce 1964; Howar
d and Oliver 1978; Lemke
and Kissam 1988; Schmidt 1984; Suitm
erlin et al. 1977,
111


104
Fig
4-10 Aleochara notula preying on horn fly eggs under
laboratory conditions.


). Mean numbers of
123
pitfall traps were operated six weeks after the Amdro
application (August 23, 1993). Th^ mean number of
staphylinids collected in the traps in the Amdro-treated
area was significantly greater than that in the control area
(F(1 18) = 15.86, P < 0.01; Table 5-4
hydrophilids, carabids, and scarabs were low in both the
Amdro-treatment and controlled area, but the mean number of
hydrophilids collected in the Amdro-treated area was
significantly greater than that in
6.71, P < 0.05; Table 5-4). Fall a
Spodoptera fruoiperda (Smith), wer
time. The mean number of fall army
significantly greater in the Amdro-
the control area (F
rmyworm larvae,
e commonly collected this
worm larvae collected was
treated area than that in
the control area (F(1 18) = 15.79, P < 0.01; Table 5-4).
Pitfall trap collections in 0
showed that the most species taxon
in pitfall traps was Staphylinidae
1992, the mean number of Philonthus
Amdro-treated area (2.7 2.93) was
(t(i8) = 2.217, P < 0.05) than those
0.5). In 1993, the mean number o
collected in the Amdro-treated area
significantly greater (t(18) = 3.55,
the control area (0.5 0.53). P.
predator of the horn fly (Fincher a
(1,18)
ctober of 1992 and 1993
of arthropods collected
(Table 5-3 and 5-5). In
spp. collected in the
significantly greater
in the control area (0.5
f Philonthus spp.
(4.40 3.43) was highly
P < 0.01) than those in
loncricornis. an effective
nd Summerlin 1994), was
common in the Amdro-treated area cojmpared with the control


113
bait Amdro (American Cyanamid Compa
ny, Wayne, NJ),
containing active ingredient--hydra
methylnon (0.73%) and
inert ingredients (99.27%). No res
idue is left in the
environment. Amdro granules are hi
ghly attractive to fire
ant workers and are carried direct!.
y into the mound by
workers as food for the colony, ki!
ling the queen and the
whole colony. The suppression of f
ire ant populations by
Amdro will continue for several mor
ths after application
(Collins et al. 1992).
In September 1992 and August 1
993, Amdro was applied on
the pasture surface by a tractor me
unted auger applicator
(Williams et al. 1983). This equip
ment consists of an auger
conveyer, a 1/3-hp electric motor a
nd gear drive, a hopper,
an electric motor and spreader, a k
ait deflector, and a
support frame. One pound Amdro per
acre was applied in the
treatment area.
Fire ant populations were moni
tored by baited traps (Oi
et al. 1994). Traps were polyethyl
ene hinge-cap vials (50
ml; 8 cm length X 3 cm diam) and fi
lied with 20 g fresh
ground beef. All ants could enter
and exit the traps until
the traps were collected. Ten trap
s were positioned every
15 m in grid pattern in the Amdro-t
reated and the control
area. If the position happened to
be on a fire ant mound,
the trap would be repositioned near
by to avoid the mound.
The traps were set in the pasture a
t 10:30 am and collected
in an hour and capped. Ants were c
ounted after being frozen


67
include house flies, flesh flies and horn flies (Wharton
1977; Sanders and Dobson 1966). Three species of Eucoilidae
Trvblioqraoha sp., Kleodotoma sp., and Rhoptromeris sp.)
were represented; three genera have all been recorded as
parasitoides of fly puparia (Krombein et al. 1979).
Gonatocerus spp. are recorded as parasitoids of homopteran
eggs (Krombein et al. 1979). Spalangia cameroni
(Pteromalidae) was reared out from the puparia of H.
irritans and Brontaea cilfera; it has been recorded as a
parasitoid of horn fly pupae by Escher (1977) and Butler et
al. (1981) in Florida. Trichopria spp. are recorded as
parasitoids of fly puparia (Krombein et al. 1979).
Telenomus spp. are recorded as parasitoids of Hemiptera and
Lepidoptera (Krombein et al. 1979) Gambrus ultimus is
recorded as a parasitoid of lepidopterous larvae and has
been found in lepidopterous cocoons (Townes and Townes
1962) .
Other Insects and Animals
Orthoptera. A cricket, Nemobius fasciatus (DeGeer),
has been reported to prey on horn fly pupae under laboratory
conditions (Bourne and Nielsson 1967). Crickets were
commonly collected in pastures in the present study and in
previous studies (Bourne and Nielsson 1967; Schoenly 1983).
Acari. Mites were commonly collected by trapping and
extracting and most of them were brought to the fresh dung


37
1991; Hunter et al. 1986b, 1989; Tawfik et al. 1976a, b, c).
The immature stages have been described for
Staphylininae (Frank 1991; Mank 1923; Hunter et al. 1989;
Tawfik 1976a, b, c; Hinton 1981a&b; Wu and Zhang 1990),
Oxytelinae (Frank 1991; Hinton 1944; Legner and Moore 1977),
Aleocharinae (Peschke and Fuldner 1977; White and Legner
1966), and other subfamilies (Frank 1991).


Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment
of the Requirements
for the Degree of Doctor of Philosophy
THE ARTHROPOD COMMUNITY IN PASTURES AND ITS BIOCONTROL
POTENTIAL FOR THE HORN FLY, Haematobia irritans (L.)
IN NORTH-CENTRAL FLORIDA
By
Guangye Hu
May 1995
Chairperson: J. Howard Frank
Major Department: Entomology and Nematology
A complete arthropod community was investigated by
pitfall traps and emergence boxes from 1991 to 1993. Over
60,000 invertebrates were collected in pastures. Arthropods
found were 226 species belonging to 73 families in 14
orders. Coleptera were the most diverse as judged by
trapping and extracting (109 species), Diptera the second
(35 species) and Hymenoptera the third (24 species). Most of
the beneficial insects were in these three orders.
Field mortality of horn flies caused by the arthropod
community was tested by seeding horn fly eggs underneath
artificial cowpats, which were either exposed or isolated
from the arthropod community by using cone traps. The
community-caused mortalities of horn flies in the field were
75.9% and 66.7% in July and August 1992, respectively. The
xi


88
Fig.
4-4. P_;_ longicornis adult pr
under laboratory conditions.
eying on horn fly
eggs


8
Egg. Most of the eggs laid are reddish and difficult
to detect in manure (Hogsette and Koehler 1986), but a small
percentage of tan, yellow, and white eggs is also laid
(Morgan and Schmidt 1966). The egg is 1.2 mm long and 0.32
mm wide (Miller et al. 1984) and is usually deposited on the
undersurface of the edge of fresh dung pats. The embryo
hatches from the egg less than 24 hours after deposition in
the field during the summer, but embryonation time depends
upon temperature (Melvin and Beck 1931; Melvin 1934; Depner
1961). Temperature and moisture conditions affect the egg
development significantly (Bruce 1964; Lancaster and Meisch
1986; Wilkerson 1974).
Larva. The horn fly larva is cream-colored, with the
anterior pointed and the posterior truncate. It has a mouth
hook at the front, a pair of 4-6 branched anterior spiracles
and a pair of heavily pigmented knob-like posterior
spiracles. The larva passes through three instars in the
manure within four days (Wilkerson 1974). The newly hatched
larvae immediately seek a crack, crevice, or perforation in
the manure into which they crawl to obtain food and shelter.
When the manure pat crust becomes firm and dry, horn fly
larvae move to a more suitable part of the manure (Bruce
1942, 1964). About 92.5 hours after hatch, larvae migrate
for pupation to the underside of the manure or into the soil
depending on the relative moisture content of the
alternatives (Bruce 1964; Escher 1977).


70
Table 3-7 (continued)
Trapped Extracted
Arachnida (Araneae) 134 ++
Linyphiidae
Eperigone banksi Ivie & Barrows +
Erigone autumnalis Emerton +
Meinoneta sp. +
Diplopoda
Polyzoniida (1 species) ++ +
Nematomorpha
Gordius sp. 3
Nematoda
Rhabiditidae
Coarctadera coarctata (Leuckart) ++
Mollusca 3
* Acari (mites) were determined by G. Krantz and M.
Hennessey; Araneae (spiders) by G. Edwards; Collembola by R.
Snider; Orthoptera, Hemiptera and Homoptera by F. Mead and
T. Henry; Lepidoptera by D. Habeck; Nematodes by K. Nguyen
and R. Esser; and Dermaptera by J. Watts.


134
inorionella (Sarcophagidae) Lestodiplosis spp.
(Cecidomyiidae), Palaeosepsis insujaris (Sepsidae), and
Forcipomyia brevipennis (Ceratopogonidae), Bradvsia
coprophili (Sciaridae), and Svlvicola notlalis
(Anisopodidae) .
Discussion
Experiments in the fall of 1992 and 1993 showed that
RIFA had a positive effect in reducing horn fly populations
emerging from the cowpats in pastures of north-central
Florida. The results agreed with those of Summerlin et al.
(1977), Schmidt (1984), Howard and Oliver (1978), and Lemke
and Kissam (1988). However, estimates of RIFA-induced horn
fly mortality have been diverse. Summerlin et al. (1977)
showed that RIFA reduced emergence of adult horn flies about
20 fold from bovine feces in a laboratory study in Texas.
Schmidt (1984) showed a sevenfold increase of horn fly
numbers in a pasture in Texas where RIFA was controlled.
Howard and Oliver (1978) found that 2-5 times more horn fly
pupae were recovered from pasture ii
was controlled by Mirex bait than ii
was not controlled. Lemke and Kissc
the number of horn flies emerged frc
RIFA was controlled using Pro-Drone was 55% greater than the
number that emerged from piles in a RIFA-infested field in
n Louisiana where RIFA
n a pasture where RIFA
am (1988) reported that
om manure piles where


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utson (eds.): Biological
versity of California,
Anderson, J.R. and E.C. Loomis. 1978. Exotic dung beetles
in pasture and range land ecosystems. Calif. Agrie. 32:
31-32.
Ashmead, E.H. 1894. A synopsis of t
America. Proc. Entomol. Soc. W
Axtell, R.C. 1963. Effect of Macroohelidae (Acaria:
he Spalangiidae of North
ash. 3: 27-37.
Mesostigmata) on housefly prod'
manure. J. Econ. Entomol. 56:
uction from dairy cattle
317-321.
Barlett, A.C.
rearing.
1984. Advances and challenges in insect
U.S. Dept. Agrie., Agrie. Res. Serv. pp 1-8.
Beadles, M.L., J.A. Miller, W.F. Ch
and R.L. Harris. 1975. The ho
drinking water of cattle for c
68: 781-785.
amberlain, J.L. Eschle,
rn fly: Methoprene in
ontrol. J. Econ. Entomol.
Blackwelder, R.E. 1943. Monograph of the West Indian beetles
of the family Staphylinidae. U. S. Natn. Mus. Bull.
182: 1-658.
146


Bulls often have larger numbers than cows. Young calves
seem to be less bothered than either cows or bulls. Horn
fly adults in Florida reach 10,000-20,000 flies per animal
with bimodal peaks in abundance in May-June and August-
September (Butler 1990).
The horn fly is one of the most damaging pests of
cattle, especially in southern areas where fly populations
may reach several thousands per animal during the long fly
season (Kinzer 1970). Damage caused by the horn fly is due
to irritation of the host, loss of blood, reduced vitality,
and refusal to graze when the numbers are high (Bruce 1964).
The repeated biting of hundreds to thousands of flies
daily irritates cattle. The energy extended in efforts to
dislodge the flies, including tail switching, head slinging,
walking and aggregation (Harvey and Launchbaugh 1982),
causes unmeasured losses. Both sexes of the adults suck
blood. Daily feeding frequencies are reported as two or a
few times (Bruce 1942, 1964) to 20 times (Koehler and Butler
1980), or even more frequently (Harris et al. 1974). Meal
size taken per fly varies from 0.045 to 0.19 mg. Feeding
occurs throughout the day. In the laboratory, Harris et al.
(1974) observed that females fed up to 38 times a day, with
an average of 163 minutes feeding per day; males fed 24
times per day, with an average of 96 minutes feeding per
day. A population of 10,000-20,000 flies per animal would
cause a loss of 2 liters of blood per month (Butler 1975).


119
Table 5-1. Numbers of horn flies ei
pats in the Amdro-treated and
1992 and 1993
njierged from artificial
control areas, October
Untreated
Treated
October 1992
Mean
Range
SD
n
October 1993
Mean
Range
SD
n
0.11
0-1
0.31
18
t = 3.44
1.08
0-4
1.17
12
t = 2.63
1.94
0-6
2.17
18
P < 0.01
2.91
1-7
0.57
12
P < 0.05


81
to the trap) to cover the hole to collect horn flies emerged
h of the vial was fitted
nt the flies from going
from the artificial pat. The moutl'
with a saran screen funnel to preve
back to the container. The cages were maintained at 27C,
50-60% RH and a 14:10 (L:D) photoperiod. On the 8th day
after the eggs were seeded, all the containers were covered
by a piece of black polyethylene through which holes (5.2
cm) were cut to expose the vials to the light. Attracted by
light, the flies went to the funnejled vial soon after they
emerged. Replicates for treatments with one beetle were 10
with P_;_ longicornis. 14 with P^ f lavolimbatus. 10 with P.
ventrales, 7 with P^. hepaticus, and 3 for control,
respectively; replicates for treatments with two beetles
were three each for P^_ longicornis. P. f lavolimbatus. and P.
hepaticus.
Differences in mean horn fly emergence between
treatments were analyzed by a one-way completely randomized
analysis of variance (ANOVA), and the significance of
differences between means was tested by Duncan's multiple
range test (SAS 1990). The numbers of adult horn flies
survived of predation were transformed by log (N+l) to
reduce heteroscedasticity (Dowdy and Wearden 1983) before
T
ANOVA was conducted.


26
(Summerlin et al. 1984b) in Texas and Staphylinidae in
Louisiana (Howard and Oliver 1978).
Mites. Several species of mites encountered in dung
are predators of flies. These mites are in the following
families: Parasitidae, Uropodidae, Eviphididae, Laelapidae,
Pachylaelapidae, and Macrochelidae (Krantz 1983).
Macrochelidae, in particular, have been considered to have
potential for biocontrol of dung-inhabiting flies (Anderson
1983; Doube et al. 1986; Krantz 1983; Axtell 1963) and have
been shown to be efficient predators of house flies (Axtell
1963; Krantz 1983), bush flies, face flies and horn flies
(Halliday and Holm 1987; Anderson 1983). These mites
commonly appear on the surface of droppings as soon as
beetles arrive, because many are mainly carried phoretically
by dung-inhabiting beetles including Scarabaeidae (Mohr
1943; Poorbaugh et al. 1968; Krantz 1983; Stewart and Davis
1967), Trogidae and Histeridae (Stewart and Davis 1967).
Halliday and Holm (1987) tested nine species of
macrochelid mites as predators of the bush fly and horn fly.
The mites preyed on fly eggs and larvae, but preferred
larvae over eggs. Macrocheles peregrinus (Krantz), M. glaber
(Mller) and M_;_ peniculatus Berlese were the most efficient.
In Australia, Macrocheles perecrrinus has been imported
from Africa and established (Roth et al. 1988b). This
species was shown to attack horn tly eggs in all stages of
development, and to kill the larvae 24 hours after hatching.


29
dung, survival from eggs to adult flies should be reduced if
dung deposits are rapidly buried or eaten by dung beetles
(Fincher 1986). There are approximately 5,000 species of
Scarabaeidae worldwide (Skidmore 1991). Certain species in
the subfamilies Aphodiinae, Scarabaeinae, and Geotrupinae
are usually abundant (Hanski 1991; Woodruff 1973; Skidmore
1991). Most scarab species belong to the subfamily
Scarabaeinae, which contains 4,00(} species (Bornemissza
1976).
Coprophagous beetles cause mortality of horn flies by
competing for the same food source (cattle dung), and by
moving and burying dung to reduce and interrupt larval
habitats of the horn fly (Anderson and Loomis 1978; Blume et
al. 1973; Bornemissza 1970, 1976; Doube and Moola 1988;
Ferrar 1975; Fincher 1981, 1986, 1990; Waterhouse 1974).
These beetles also reduce dung accumulation, improve
pastures by increased fertility and improved soil structure
(Fincher 1981, 1990; Fincher et al. 1983). Rapid removal
of feces would return areas of pasture to grazing (which
normally would be lost because of
tons of nitrogen normally lost into the atmosphere, and
reduce pest fly populations on livestock. Dung tunnelers
also provide runways so predatory staphylinids can get at
the flies (Valiela 1974). The larvae of dung beetles alsc
digest bacterial albumens which may account for their
subsistence in old dung heaps (Merritt 1976). The benefit
contamination), recycle


48
emergence boxes. These arthropods either were in the adult
stage when the dung was collected, or had developed from
immature stages within the dung.
Insect diel activity was measured by pitfall traps
baited with fresh dung. Fifteen traps were run continuously
for 24 hours in pasture A in July and August 1992,
respectively. At each 6-hour interval, the bait in the
traps was replaced with fresh dung and the trapped insects
were collected.
Results and Discussion
Over the sampling period from 1991 to 1993, more than
50,000 invertebrates (Collembola were not included) were
collected in two pastures by two collecting methods. In
total, 226 species of invertebrates were collected and
identified. Arthropods found were 223 species belonging to
79 families in 14 orders. Coleptera were ranked first in
number of species (109), Diptera second (35), and
Hymenoptera third (24) (Table 3-1).
Coleptera
Staphylinidae held the most species (44) (Table 3-2),
followed by Scarabaeidae (27) and Carabidae (20). Four
species each of Histeridae and Hydrophilidae were collected
(Table 3-4). Staphylinidae accounted for the greatest


Dollar losses to the cattle
6
mdustry due to horn flies
are increasing. The annual losses
3 were estimated as three
million in 1942 (Bruce 1942), 150
million in 1956 (Knipling
and McDuffie 1956), 179 million ii
i 1976 (Steelman 1976), 730
million in 1981 (Drummond et al.
L981), and 870 million in
1991 (Kunz et al. 1991), respectively. In Florida, Koehler
and Butler estimated losses were
?36 million in 1976 and $50
million in 1984, respectively; Butler later estimated a
loss of about $61 million in 1986
(Hogsette and Koehler
1986).
The losses include the reduction of weight gain, or
weight loss in cattle (Cheng 1958
; Cutkomp and Harvey 1958;
Roberts and Pund 1974; Campbell 1976; Harvey and Brethour
1979; Laake 1946), as well as reduction in milk production
in dairy cattle (Bruce and Decker
1947; Granett and Hansens
1956, 1957). Weight gains of 30
to 70 pounds were observed
after animals were treated with DDT (Laake 1946). Grannett
and Hansens (1957) showed a decrease in milk production of
one-fourth in unprotected dairy herds. Bruce (1947) showed
a high inverse correlation betweei
n changes in milk
production and fly abundance.
The damage threshold of fly
numbers per cow was
reported to be from 50 (Butler 1975) to 200 (Hogsette et al.
1991, Schreiber et al. 1987b), or
even higher (Haufe 1982).
The heterogeneity of these data i
s caused by the
experimental design, the region w]
tere the tests were


17
America. Staphylinidae, for example, have been considered
the most important predators in cattle dung. Fourty-three
species were found to be associated with cattle dung in this
country (Fincher 1990), while 133 species were reported from
cattle dung in a study in Finland (Koskela 1972). The horn
fly is also an immigrant pest in Australia and has plagued
the cattle industry there, but it rarely reaches pest status
in Africa (Zumpt 1973), because there is a higher level of
fauna-induced mortality in Africa than in Australia
(Bornemissza 1960; Fay 1986). Natural enemies inhabiting
cattle dung in Europe, Africa and Asia have been introduced
for horn fly control in North America (Fincher 1990; Merritt
and Anderson 1977). Introduction of additional biocontrol
agents for biocontrol of the horn fly is needed (Fincher and
Morgan 1990; Fincher 1990; Fincher and Summerlin 1994),
because arthropods currently inhabiting cattle dung have not
suppressed the horn fly populations to an acceptable level
in North America (Fincher 1990; Hunter et al. 1991).
Dung Arthropod Community
Dung Arthropod Community Composition
Basic information on the arthropod fauna of dung is
essential for consideration of introduction of biocontrol
agents for horn fly control. Bovine manure supports a large
population of insects, mainly including many dipterous,


40
as a model for
replaced with a piece of
pats about 24 hours old were collected from pasture A and
taken to the laboratory to extract the dung-associated
arthropods. Five dung pats were sampled twice a month from
May to October and once a month from November to April.
Dung pats were also sampled from pasture B in October 1992
and 1993. The pats were placed individually into emergence
boxes of a type used by G. T. Fincher (USDA-ARS, College
Station, Texas) who provided one,
construction of others. Each box was a gray plastic kitchen
box 46 cm long X 33 cm wide x 18 cm high. A 30 x 20 cm2
section was cut from each lid and
black cotton cloth to provide ventilation. A circular hole
(4 cm diameter) was cut through one end of the box. The lid
for a 7.5 cm high X 4 cm diam vial was perforated, then
glued and riveted to the box over the hole. When the vial
was screwed onto the box over the hole, it served as a
collection device to collect arthropods that attempted to
escape the emergence box by flight. In a similar manner, a
10 cm diam hole was cut through the bottom, and the lid of a
12.7 cm deep X 12.7 cm diam plastic cup was perforated, and
glued and rivetted over the hole and then the cup was
screwed onto the lid. This device
walking and falling into it. The
cup were each fitted with a hardware cloth funnel to prevent
insects from escaping back into the box. The vial and the
cup collected adult arthropods that left the dung in the
collected arthropods
mouths of the vial and the


10
piercing and sucking mouthparts and are blood feeders.
Mating and Oviposition
Mating
Adults began to mate one day after eclosin under
natural conditions (Harris et al. 1968), but two or three
days after eclosin under laboratory conditions (Bruce 1964,
Lancaster and Meisch 1986). Females are monogamous, but
males can inseminate an average of 4.6 females in the field
(Harris et al. 1968).
Mating behavior has been described by Bolton (1980) and
Zorka and Bay (1980). Mating behavior of the male involves
orientation, tapping the dorsum of the female's abdomen with
prothoracic tarsi, mounting and using its legs to grasp the
female, positioning genitalia towards the female's, grabbing
the female's ovipositor with claspers, and copulation.
Copulation lasts from 0.9 to 5 minutes (Zorka and Bay 1980),
3.5 to 11.5 minutes (Bolton 1980), and 0.5 to 5 minutes
(Bruce 1964). Mating is chemically mediated and males are
attracted by cuticular lipids of the female (Mackley et al.
1981).
Oviposition
Females begin laying eggs one day after mating (Bruce
1942, 1964; Harris et al. 1968; Schmidt et al. 1972) and
oviposit at any time during the day and night (Sanders and
Dobson 1969; Kunz et al. 1970). Horn fly eggs are deposited


19
dung in the United States.
Mohr (1943) reported 67 species of insects inhabiting
dung, representing 28 families of
5 orders in Illinois.
Poorbaugh et al. (1968) reported
.51 species of insects
attracted to and reared from cowj
3ats in California,
representing 76 families in 4 orders. Sanders and Dobson
(1966) reported 38 insect species
including 17 families of 3
orders in Indiana. Blume (1970) i
reported 103 insect
species representing 45 families of 5 orders in Texas.
Macqueen and Beirne (1975) reported 67 arthropod species
representing 21 families of 4 orders in British Columbia,
Canada. Valiela (1969b) reported
109 species of insects and
mites in 43 families of 7 orders
m New York. Cervenka and
Moon (1991) reported 108 arthropoc
1 species in 19 families of
4 orders in Minnesota. Wingo et al. (1974) reported 157
arthropod species in 33 families
of 5 orders in Missouri.
Succession of the Community
Members of the newly forming
dung arthropod community
start to arrive immediately, with
the horn fly prominent
among the first colonists. Many c
)ther species arrive during
the first few minutes such as Sarcophagidae (Mohr 1943). And
the major period of colonization 1
asts until a distinct
crust of the dung has formed (Skic
Imore 1991). Hunter et al.
(1986a) observed that arrival time
of Staphylinidae ranged
from 30 minutes to 24 hours. The
insects arriving within


31
1990). Hence, further introductior
i of dung beetles is needed
in this country.
Fullaway (1921) introduced dung beetles in Hawaii as a
biological control agent for the horn fly. Twenty-three
species of dung beetles were introduced in Hawaii from 1906-
1963 (Fincher 1986). During the 1970s, several additional
species were released in Hawaii b^
courtesy of the CSIRO
Dung Beetle Project from Australia
Experiments in Hawaii
in 1966 showed 95% fewer horn flies emerged from cowpats
attacked by Onthophaqus gazella than from pats from which
these beetles were excluded (Bornemissza 1970). So far, 15
species of dung beetles have been released in the
continental USA (Fincher 1990). The following species have
been established: O. taurus Schreber, which was released in
Texas and established in southern States (Fincher and
Woodruff 1975; Fincher 1990) and California (Anderson and
Loomis 1978); 0. gazella, which was released in Texas
(Blume and Aga 1978), California a
nd Georgia in 1975,
Arkansas in 1976 and Mississippi i
n 1979 (Fincher 1981) and
established in southern States (Fi
ncher et al. 1983; Hunter
and Fincher 1985); 0. deoressus Ha
rold which was established
in Florida and Georgia by unknown means (Fincher 1990);
Euoniticellus intermedius Reiche.
which was released and
established in Texas (Blume 1984);
Onitis alexis Klug,
which was released in Texas and es
tablished in California
(Anderson and Loomis 1978).


76
those for laboratory
(5 cm diameter) was cut
along the outside of the fence surrounding plots in the
pastures. Adjacent pats were separated by 10 m. Egg-seeding
procedures (100 eggs/pat) followed
survival studies. Ten pats (odd numbers) were at first open
to allow other insects to come to the dung and then covered
by cone traps on the 8th day after egg-seeding; the other 10
(even numbers) were covered immediately after egg-seeding.
The cone traps (Fig. 4-1) used for covering the seeded
cowpats were constructed of a wire frame and wrapped with
fine saran screen. The trap was 30.5 cm diameter (bottom)
and 50.8 cm high. A circular hole
through the screen on the top of the trap, and a 8.89 cm
high X 5.08 cm diameter vial was screwed on (the lid of the
vial was perforated, then glued and rivetted to the trap) to
cover the hole to collect horn flies that emerged from the
pat. The mouth of the vial was fitted with a saran screen
funnel to prevent insects from escaping back to the trap.
The vials were checked daily from the 8th day after egg-
seeding until two days after the last horn flies were found.
Differences in mean numbers of horn flies emerged
between treatments were analyzed by a Student's t-test
(SigmaPlot 1994). Before the t-test was conducted, the
numbers of the horn flies were transformed by log (N+l) to
satisfy equal variance and normality assumpted by t-test.


ACKNOWLEDGMENTS
I wish to express my deepest gratitude to my major
professor, Dr. J. Howard Frank, for his invaluable guidance,
encouragement, advice and financial support. Thanks are
also extended to Drs. J. F. Butler, L. P. Lounibos, J. A.
Hogsette, R. S. Sand for serving on the supervisory
committee and contributing to the completion of the
dissertation. I wish to extend special thanks to Dr. Butler
for his help with photographing insects, allowing me to use
the equipment at his laboratory, and providing me access to
his horn fly colony for my experiments, and Dr. Lounibos for
loaning me a video camera for my research.
I am also grateful to Mr. P. Dixon and J. Stokes for
granting me use of the beef pastures of IFAS, University of
Florida, for my field research. I wish to acknowledge Dr.
D. Williams for the help with application of Amdro for fire
ant population control, Dr. T. Fincher for donating an
emergence box designed to extract insects from cowpats, and
Dr. J. Castner for photographing emergence and testing
equipment used for this study.
I would like to thank the following specialists for
helping me with the insect identification: K. Ahlmark, F.
ii


16
1938), respectively. An updated version is being tested as
the University of Maryland (Hogsette, pers. comm.).
Less attractive hosts, such as Brahman (zebu) or
Brahman-cross cattle (Tugwell et al. 1969; Steelman et al.
1994) have been reported to decrease horn fly populations.
However, weight gain with these cattle are usually less than
those of English breeds (Steelman et al. 1994).
Studies show that exposure of animals to hematophagous
arthropod ectoparasites, such as ticks, mosquitoes and horn
flies, evokes an immune response to the parasite (Kerlin and
Allingham 1992). Developing a vaccine would enhance the
host immune response which would be deleterious to
arthropods' feeding on the host. Vaccines are possibly
developed to inoculate cattle against horn fly bites in the
future.
Biological Control
Though research emphasis on horn fly control in North
America has been centered on using chemical insecticides,
increasing attention has been given in the past 2-3 decades
to an examination of the field relationships and
interactions between immature stages of the horn fly and the
other organisms that share its habitat.
The horn fly is a serious pest in North America, but
not in its origin, Europe, because the insect fauna of
droppings is reportedly richer in Europe than in North


93
ventralis. and P^_ flavolimbatus.
The results of this study suggest that all the four
species are effective predators. They were often collected
by trapping and extraction during the survey. P,
lonqicornis adults (6.0-7.5 mm), larger than those of the
other species (4.5-5.5 mm) tested in the present study, have
a higher predation rate than do Ft. f lavolimbatus. P.
ventralis and Ft. hepaticus. P. lonqicornis. of cosmopolitan
distribution (Blackwelder 1943), is well-established in
north central Florida and may be the most effective
predator.
Fincher and Summerlin (1994) compared Ft. loncricornis
with Pi flavocinctus and Ft. minutus under simulated field
conditions. Their results showed that It. lonqicornis had a
higher predation rate than those of the other two species,
whose size (4.5-5.5 mm) is similar to that of P.
f lavolimbatus. P. hepaticus and It. ventralis. The results
of Fincher and Summerlin (1994) are similar to those of the
present study.
Predation by P. lonqicornis at varying prey densities.
When one Pt_ lonqicornis was maintained per 100 g manure with
varying numbers of horn fly eggs, the number of the horn
flies destroyed increased in relation to egg density (Fig.
4-8). The result shows that Ft. lonqicornis will increase
its efficiency when horn fly numbers increase in the manure.
Fincher and Summerlin (1994) showed that Ft. lonqicornis were


I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
Robert S. Sand
Associate Professor of
Animal Science
This dissertation was submitted to the Graduate Faculty
of the College of Agriculture and to the Graduate School and
was accepted as partial fulfillment of tl
the degree of Doctor of Philosophy,.
May 1995
,e requirements for
/? 'dt -
Dean, College of
Agriculture
Dean, Graduate School


116
RIFA/trap were collected in the Amc
ro-treated area and the
control area, respectively. The me
an number in the control
area was significantly greater thar
that in the Amdro-
treated area (t(18) = 5.14, P < 0.01
Fire ant numbers were
80% less in the Amdro-treated area
compared with the control
area. Three weeks after the applic
ation of Amdro, an
average of 5.7 15.1 RIFA/trap was
collected in baited
traps from the Amdro-treated area,
while an average of 184
125 fire ants/trap was collected fr
om the control area. The
difference was highly significant (
t(18) = 4.43, P < 0.01).
Fire ant numbers were 95% less in t
he Amdro-treated area
compared to the control area. Four
weeks after the
application of Amdro, horn fly eggs
were seeded in the
artificial pats in the pastures and
the epigean arthropods
were collected (October 28).
In 1993, Amdro was applied on
September 3 when baited
traps showed there was no significa
nt difference between the
mean fire ant numbers collected frc
m the Amdro-treated and
the control areas. Four weeks late
r, the mean number of
fire ants collected from the contrc
1 area (214 154) was
significantly greater (t(18) = 4.0,
? < 0.01) than that in
the Amdro-treated area (18 53).
Though fire ant numbers
were over 90% in the Amdro-treated
area compared with the
control area, 47 small new fire ant
mounds were found in the
Amdro-treated area. This is becaus
e RIFA has a multiple-
gueen system and splits into multip
>le mounds quickly when


Table Page
5-3. Numbers of staphylinid specimens collected by
pitfall traps from the Amdro-treated and control
areas in October 1993 121
5-4. Numbers of arthropod specimens collected by pitfall
traps from the Amdro-treated and control areas in
October 1993 125
5-5. Numbers of staphylinid specimens collected by
pitfall traps from the Amdro-treated and control
areas in October 1993 126
5-6. Numbers of specimens of arthropod taxa collected
per cowpat sample from the Amdro-treated and
control areas, October 1992 129
5-7. Numbers of specimens of selected arthropod taxa
collected per cowpat sample from the Amdro-treated
area and the control area, October 1993 131
5-8. Numbers of staphylinid specimens extracted from
Amdro treatment and control area in October 1993 133
vm


and all of the predatory beetles used for the predation
tests were extracted from cow-dung
hours. Therefore, predation plays
suppressing horn fly populations in
deposited within 24
an important role in
pastures in north
central Florida.


I certify that I have read this study and that in my
opinion it conforms to acceptable standards of scholarly
presentation and is fully adequate, in scope and quality, as
a dissertation for the degree of Doctor of Philosophy.
I certify that I have read thi
opinion it conforms to acceptable
presentation and is fully adequate
a dissertation for the degree of D
J. Howard Frank, Chair
Professor of Entomology
and Nematology
s study and that in my
standards of scholarly
in scope and quality, as
ctor of Philosophy.
I certify that I have read thi
opinion it conforms to acceptable
presentation and is fully adequate
a dissertation for the degree of Do
I certify that I have read thi
opinion it conforms to acceptable
presentation and is fully adequate
a dissertation for the degree of D
F. Butler
Professor of Entomology
and Nematology
s study and that in my
tandards of scholarly
in scope and quality, as
ctor of Philosophy.
L. Philip Lounibos
Professor of Entomology
and Nematology
s study and that in my
standards of scholarly
in scope and quality, as
octor of Philosot
3¡y-
[erme A. Hog
Assistant Professor
Entomology and
Nematology


Table 3-4. Coleptera collected in pastures in north central
Florida, with exclusion of Staphylinidae and
Scarabaeidae. The occurrence codes for
enumerated: (+) rare, (++) common, and
species not
(+++) abundant.
Trapped
Extracted
Carabidae
Tachys sp.
36
++
Panagaeus fasciatus (Say)
1
Evarthrus morio Dejean
3
Calosoma sayi Dejean
6
Ardistomis viridis Say
22
+
A. puncticollis Putzeys
9
+
Dyschirius sp.
2
Clivina sp.
2
Aspidoglossa subangulata Chaudoir
7
Stenolophus infascatus (Dejean)
1
Selenophorus paliatus F.
9
+
S. discopunctatus Putzeys
2
S. fossulatus Dejean
4
Amblygnathus irepennis (Say)
1
Amara sp.
1
Stenocrepis quatuordecimstriata
36
++
Chaudoir
Galerita sp.
1
Tetragonoderus intersectus
20
++
Haldeman
Pasimachus sublaevis Dejean
8
P. marginatus (F.)
2
Histeridae
Hister coenosus Erichson
26
11
Phelister haemorrhous Marseul
5
2
Acritus ignobilis Lewis
22
7
Saprinus pennsylvanicus (Paykull)
28
6
Hydrophilidae
85
Sphaeridium lunatum (L.)
34
5
Cryptopleurum subtile Sharp
3
Cercyon variegatus Sharp
199
42
C. atricapillus (Marsham)
134
31
Tenebrionidae
Gondwanocrypticus obsoletus(Say)
13
4
Poecilocrypticus formicophilus
2
4
Gebien


27
Each mite has the capacity to kill 8-10 flies when there is
a high density of eggs provided. It caused 66% and 54%
suppression of horn fly populations, respectively, in two
field tests (Doube et al. 1986). In another experiment,
however, M. pereqrinus caused an average of 33% suppression
of horn flies in field cowpats, and had a stronger
preference for eggs of other dipterous species that have
softer chorions (Roth et al. 1988b).
Flies. Although some 200 species of Diptera have been
reported on or in cattle dung, on]
in the dungpats can contribute to
flies by competition and predation. Members of the families
Muscidae, Empididae and Sarcophagidae have been considered
to be possible predators (Harris and Blume 1986). The flies
reported to be predacious are Drapetis spp. (Empididae)
(Laurence 1952), Hvdrotaea (Muscidae) (Merritt 1976; Hammer
1941), Myospiia meditabunda (F.)
al. 1968), Gvmnodia f-Brontaea) spp. and Orthelia spp.
(Muscidae) (Ferrar 1975) and Ravinia Iherminieri (Robineau-
y the flies that develop
the reduction of the pest
(Muscidae) (Poorbough et
Desvoidy) (Sarcophagidae) (Pickens 1981). Hammer (1941)
reported that Hvdrotaea larvae are typically facultative
predators. Data on suppression of horn fly populations by
these species are unavailable. Mvospila meditabunda
(Muscidae) was assumed to inflict high mortality on horn fl
and other coprophagous larvae (e.g. sarcophagids,
scatophagids, sepsids, and sphaerocerids) in California


Table 4-1. Larval predation rates c
during the whole developmental
and larvae under laboratory cc
91
Df five Philonthus species
L period on horn fly eggs
Dnditions.
Prey
Species stages
No.
predators
Mean
SD
Range
P. loncricornis
eggs
33
196 .'
3 60.94
84-369
larvae
33
125 .:
L 35.13
70-233
P. flavolimbatus
eggs
22
47.1
11.54
27-65
larvae
22
59.5
16.85
24-86
P. sericans
eggs
35
36.3
25.19
32-12
larvae
35
44.8
18.0
22-104
P. heoaticus
eggs
6
20.7
6.82
7-26
larvae
6
30.0
9.78
14-42
P. ventralis
eggs
29
28.9
43.56
4-195
larvae
29
142 .;
? 24.39
85-211


78
was covered by a piece of microscope slide. Oviposition and
development of the predators at different stages were
recorded daily.
Predators used in this study included 22 staphylinid
species, two hydrophilid species, one histerid species, two
carabid species, one anthicid species, and one tenebrionid
species.
Predation rate test under simulated field conditions
Predation tests for Philonthus species were conducted
in testing cages (Fig. 4-2), which are made of 15.24 cm
diameter X 5 cm high plastic containers. Moist, sandy loam
soil was placed in each container (1 cm deep). Horn fly
eggs (100) suspended in well water were pipetted on a piece
of doughnut-shaped paper towel (13 cm diam), with a circular
hole (9 cm diam) cut in the center. The paper towel was
placed on the loam soil and covered with a 100 g cattle
dung. This arrangement allowed the fly eggs to be covered
by the edge of the artificial pat, thereby simulating
natural horn fly oviposition behavior (McLintock and Depner
1954). Immediately after the eggs were seeded, one or two
adult Philonthus (mixed sexes) of each species were added to
the pat through a circular hole (2.5 cm diameter) on the
lid. Another circular hole was cut through the lid of the
container and an 8.89 cm high X 5.08 cm diameter vial was
screwed on (the lid of the vial was perforated and rivetted


127
Among the staphylinids extracted from the cowpats, only
Oxytelus incisus. and Anotylus insjqnitus were common. Few
Philonthus were extracted from the cowpats. The low numbers
of Staphylinidae compared with 1993 might be attributed to
dry weather, which caused high mortality of immature stages,
especially of Philonthus.
Other than horn flies, the mean number of the remaining
muscids from the cowpats in the Amdro-treated area (3.9
4.74) was significantly greater (F(1
Table 5-6) than that in the control area (0.6 1.8). The
mean number of sarcophagids collected from the cowpats in
the Amdro-treated area (3.6 5.21)
greater (F(1 34) = 4.53 P < 0.05; Tal
1,34)
control area (1.6 4.23). Mean numbers of small dung flies
8.5, P < 0.01;
was significantly
die 5-6) than that in the
including sphaerocerids, cecidomyii
anisopodids, and ceratopogonids col
the Amdro-treated area and the cont
significantly different (P > 0.05).
2.8 4.35 psychodids were collecte
Amdro-treated area compared with no
Mean numbers of combined hymenopter
pats in the Amdro-treated and the c
significantly different (F(1 34) = 1.15, P > 0.05).
1993 trial
Staphylinidae were more abunda
ds, sciarids,
lected from cowpats in
rol area were not
However, an average of
d from the pats in the
ne in the control area,
ans collected from the
ontrol area were not
nt in this year than in


168
Wall, R. and L. Strong. 1987. Envir
treating cattle with the antip
Nature 327: 418-421.
onmental consequences of
arasitic drug ivermectin.
Wang, S.Q., W.Z. Zhang, A.X. Xiao a
Predatory insects of fly larva
efficiency. Pest Contr. 6: 15-
nd G.Y. Zhang. 1990.
e and their predatory
18 .
Waterhouse, D.F. 1974. The biologic
Am. 230: 100-109.
al control of dung. Sci.
Watts, K.J. and R.L. Combs, Jr. 197
Haematobia irritans and other
feces in northeast Mississippi
823-826.
7. Parasites of
flies breeding in bovine
. Environ. Entomol. 6:
Wharton, RA. 1977. New world sped
Braconidae: Allysiinae), with
terminology used in the Tribe
Am. 70: 782-803.
es (Hymenoptera:
a discussion of
Alysini. Ann. Entomol.
Wharton, R.A. 1979. Some predators
breeding Diptera from central
Entomol. 55: 181-186.
and parasitoids of dung-
California. Pan-Pacific
White, E.B. and E.F. Legner. 1966.
of Aleochara taeniata, a staph
house flv. Musca domestica. An
573-577.
Notes on the life history
ylinid parasite of the
n. Entomol. Soc. Am. 59:
Wilkerson, G.G. 1974. A population
Haematobia irritans (Linnaeus)
north central Florida. Thesis,
Gainesville. 109 pp.
model for the horn fly,
(Diptera: Muscidae) in
University of Florida,
Williams, D.F., C.S. Lofgren, J.K.
1983. Auger-applicator for app
granular pesticides. J. Econ.
Plumley and D.M. Hicks,
lying small amounts of
Entomol. 76: 395-397.
Williams, D.F., C.S. Lofgren. 1983.
(Hymenoptera: Formicidae) cont
several chemicals for individu
Econ. Entomol. 76: 1201-1205.
Imported fire ant
rol: evaluation of
al mound treatment. J.
Wingo, C.W., G.D. Thomas, G.N. Ciar
Succession and abundance of in
Relationship to face fly survi
Am. 67: 386-390.
k and C.E. Morgan. 1974.
sects in pasture manure:
val. Ann. Entomol. Soc.


72
predators.
materials and fungi and are often numerous in older dung.
They mostly belong to the soil fauna (Skidmore 1991). They
also provide food for small insect
Hemiptera. Hemiptera were collected frequently by
trapping and occasionally by extraction. Many species in
this order are considered to be casual visitors to dung.
Species in the family Lygaeidae have been found to prey on
other insects in the dung (Skidmore 1991).
Hematomorpha and Nematoda. Gordius sp. (horse hair
worm) is a parasite of Coleptera, Dermaptera, Diptera,
Hemiptera, and Hymenoptera. It was recorded from the German
cockroach and Florida wood cockroach in Florida (Esser
1980). Coarctadera coarctata parasitizes staphylinids,
hydrophilids, and scarabs (Frank 1982) .
The remaining groups of miscellaneous arthropods and
other animals collected have been recorded infrequently in
the past, and their interactions with flies are poorly
known.


South Carolina. In the present study, in which the
experiments were conducted in the same pastures, horn fly
mortalities from fire ants differed between 1992 and 1993.
Heavy rain and cold weather were encountered when the
experiment was conducted in October 1993, and these factors
may have decreased horn fly numbers compared with the
experiment in 1992. Weather conditions, such as rain, dew,
soil moisture, and soil temperature, are reported to affect
foraging behavior of the ant (Lofgren et al. 1964; Porter
and Tschinkel 1987).
Results of these studies also showed positive effects
of S. invicta in reducing numbers of other muscid flies,
especially Brontaea debilis and cilfera. These flies
were reported to increase by 3-fold in a RIFA excluded area
compared to a RIFA infested area in Texas (Schmidt 1984).
Similar to the study by Howard and Oliver (1978), the
present study showed that abundance of Sarcophagidae was
reduced in an S_j_ invicta infested area compared with an area
the RIFA was controlled. Common sarcophagid species
collected in the present study included Ravinia floridensis,
R. derelicta and H. moronella. S. invicta was also shown to
reduce numbers of other Diptera, such as Sepsidae, and
Psychodidae.
RIFA was showed to adversely affect scarabaeid beetles
in the study by Summerlin et al. 1984b, even though the
effect was slight in a previous study (Summerlin et al.


Table 5-7. Numbers of specimens of
collected per cowpat sample fr
and the control area, October
131
selected arthropod taxa
om the Amdro-treated area
1993 (n = 12).
Arthropod taxa
No. collected
Amdro
Control
Staphylinidae
Mean
226.8
93.0
SD
81.89
55.16
Range
74-309
6-172
F = 4.86,
P
<
0.05
Hydrophilidae
Mean
10.17
6.5
SD
2.03
4.92
Range
6-12
0-13
F = 2.01,
P
<
0.05
Muscidae (except H_
. irritans)
Mean
15.83
4.92
SD
7.71
3.86
Range
4-33
0-12
F = 4.51,
P
<
0.10
Sarcophagidae
Mean
4.92
0.75
SD
5.51
1.01
Range
1-19
1-3
F = 4.53,
P
<
0.10
Sepsidae
Mean
44.08
28.67
SD
14.0
17.28
Range
26-74
8-60
F = 8.02,
P
<
0.05
Sphaeroceridae
Mean
19.0
39.25
SD
13.32
8.31
Range
3-4 5
0-200
F = 0.16,
P
>
0.10
Cecidomyiidae
Mean
139.83
98.42
SD
129.71
124.13
Range
7-426
0-372
F = 0.39,
P
>
0.10