Title: Florida Entomologist
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Title: Florida Entomologist
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Creator: Florida Entomological Society
Publisher: Florida Entomological Society
Place of Publication: Winter Haven, Fla.
Publication Date: 1993
Copyright Date: 1917
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Subject: Florida Entomological Society
Entomology -- Periodicals
Insects -- Florida
Insects -- Florida -- Periodicals
Insects -- Periodicals
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General Note: Eigenfactor: Florida Entomologist: http://www.bioone.org/doi/full/10.1653/024.092.0401
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Volume ID: VID00058
Source Institution: University of Florida
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(ISSN 0015-4040)


FLORIDA ENTOMOLOGIST

(An International Journal for the Americas)

Volume 76, No. 1 March, 1993

TABLE OF CONTENTS


Announcement 75th Annual Meeting and Call for Papers ............................. i


SYMPOSIUM: INSECT BEHAVIORAL ECOLOGY-'92

FRANK, J. H., AND E. D. McCoY-Introduction To Behavioral Ecology of Intro-
duction. The Introduction of Insects Into Florida ............................... 1
BENNETT, F. D.-Do Introduced Parasitoids Displace Native Ones? .............. 54
OI, D. H., AND R. M. PEREIRA-Ant Behavior and Microbial Pathogens
(Hymenoptera: Formicidae) ............................................................ 63
PARKMAN, J. P., AND J. H. FRANK-Use of a Sound Trap to Inoculate
Steinernema scapterisci (Rhabditida: Steinernematidae) Into Pest Mole
Cricket Populations (Orthoptera: Gryllotalpidae) ............................... 75
JANSSON, R. K.-Introduction of Exotic Entomopathogenic Nematodes (Rhab-
ditida: Heterorhabditidae and Steinernematidae) for Biological Control of
Insects: Potential and Problems ...................................................... 82
HABECK, M. H., S. B. LOVEJOY, AND J. G. LEE-When Does Investing in Class-
ical Biological Control Research Make Common Sense? ...................... 96
ALLEN, J. C., J. L. FOLTZ, W. N. DIXON, A. M. LIEBHOLD, J. J. COLBERT, J.
REGNIERE, D. R. GRAY, J. W. WILDER, AND I. CHRISTIE-Will the
Gypsy Moth Become a Pest in Florida? ............................................ 102


Research Reports

SHUSTER, J. C.-Xylopassaloides (Coleoptera: Passalidae) In Mesoamerica: Re-
lations, Distribution, and New Species ............................................. 114
NATWICK, E. T., AND F. F. LAEMMLEN-Protection from Phytophagous Insects
and Virus Vectors in Honeydew Melons Using Row Covers ................. 120
EVANS, G. A., H. L. CROMROY, AND R. OCHOA-The Tenuipalpidae of Hon-
duras (Tenuipalpidae: Acari) .......................................................... 126
HALL, D. G., AND R. H. CHERRY-Effect of Temperature in Flooding to Control
the Wireworm Melanotus communis (Coleoptera: Elateridae) ............... 155
BUTLER, G. D., JR., T. J. HENNEBERRY, P. A. STANSLEY, AND D. J.. SCHUS-
TER-Insecticidal Effects of Selected Soaps, Oils and Detergents on the
Sweetpotato Whitefly: (Homoptera: Aleyrodidae) ............................... 161


Scientific Notes

BREENE, R. G., D. A. DEAN, AND R. L. MEAGHER, JR. -Spiders and
Ants of Texas Citrus Groves ................................................... 168


Continued on Back Cover

Published by The Florida Entomological Society








FLORIDA ENTOMOLOGICAL SOCIETY
OFFICERS FOR 1992-93
President ............................................................................... D F. W illiam s
President-E lect .............................................................................. J. E Pefia
Vice-President ............................................................................. E .' M Thom s
Secretary ....................................................................................... D G H all
Treasurer ................................................................................... A C. Knapp
Other Members of the Executive Committee
J. L. Knapp D. P. Wojcik L. A. Wood J. Hogsette
C. S. Lofgren 0. Liburd D. R. Suiter
PUBLICATIONS COMMITTEE
C. S. Lofgren, Gainesville, FL ......................................... ................... Editor
Associate Editors
Agricultural, Extension, & Regulatory Entomology
James R. Brown-Disease Vector Ecology & Control Center, NAS, Jacksonville, FL
Richard K. Jansson-Tropical Research & Education Center, Homestead, FL
Geof Zehnder, Auburn University, Auburn, AL
Apiculture
Stephen B. Bambara-North Carolina State University, Raleigh, NC
Biological Control & Pathology
Ronald M. Weseloh-Connecticut Agricultural Experiment Sta., New Haven, CT
Book Reviews
J. Howard Frank-University of Florida, Gainesville
Chemical Ecology, Physiology, Biochemistry
Louis B. Bjostad-Colorado State University, Fort Collins, CO
Ecology & Behavior
John H. Brower-Stored Product Insects Research Laboratory, Savannah, GA
Forum & Symposia


Genetics & Molecular Biology
Sudhir K. Narang-Bioscience Research Laboratory, Fargo, ND
Medical & Veterinary Entomology
Arshad Ali-Central Florida Research & Education Center, Sanford, FL
Resumen
J. E. Pefia-Tropical Research & Education Center, Homestead, FL
Systematics, Morphology, and Evolution
Michael D. Hubbard-Florida A&M University, Tallahassee
Gary J. Steck-Florida State Collection of Arthropods, Gainesville
Willis W. Wirth-Florida State Collection of Arthropods
Business M manager ....................................................................... A. C. Knapp
FLORIDA ENTOMOLOGIST is issued quarterly-March, June, September, and De-
cember. Subscription price to non-members is $30 per year in advance, $7.50 per copy;
institutional rate is $30 per year. Membership in the Florida Entomological Society,
including subscription to Florida Entomologist, is $25 per year for regular membership
and $10 per year for students.
Inquiries regarding membership and subscriptions should be addressed to the Busi-
ness Manager, P. O. Box 7326, Winter Haven, FL 33883-7326.
Florida Entomologist is entered as second class matter at the Post Office in DeLeon
Springs and in Winter Haven, FL.
Manuscripts from all areas of the discipline of entomology are accepted for consider-
ation. At least one author must be a member of the Florida Entomological Society.
Please consult "Instructions to Authors" on the inside back cover.
This issue mailed March 31, 1993





















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ANNOUNCEMENT 76TH ANNUAL MEETING
FLORIDA ENTOMOLOGICAL SOCIETY

The 76th annual meeting of the Florida Entomological Society will be held August
9-12, 1993 at the South Seas Plantation Resort and Yacht Harbor, Captiva Island,
Florida 33924 (813-472-5111) (FAX 813-472-7541). Registration forms and information
will be mailed to members and will appear in the Newsletter.

CALL FOR PAPERS

The deadline for submission of papers and posters for the 76th annual meeting of
the Florida Entomological Society is Monday May 10, 1993. The meeting format will
be similar to that in the past with eight minutes allotted for presentation of oral papers
(with 2 minutes for discussion) and separate sessions for members presenting a poster
exhibit. There will be student paper and poster sessions with awards as in previous
years. A description of the format for judging the student paper and poster sessions will
be printed in the newsletter. Students participating in the judged sessions must be
members of the Society and registered for the meeting. Inquiries should be directed to:

Ellen Thoms
Program Committee, FES
DowElanco
One MetroCenter
Suite 240, 4010 Boy Scout Blvd.
Tampa, FL 33607
(813) 874-1200
813-877-1326 FAX










Insect Behavioral Ecology-'92: Frank & McCoy 1

Introduction To
THE BEHAVIORAL ECOLOGY OF INTRODUCTION

THE INTRODUCTION OF INSECTS INTO FLORIDA

J. Howard Frank' and Earl D. McCoy2
'Entomology and Nematology Department
University of Florida
P.O. Box 110620, Gainesville, Florida 32611-0620

'Department of Biology and Center for Urban Ecology
University of South Florida
Tampa, Florida 33620-5150

ABSTRACT

About 351 insect species have been introduced into Florida for potential release
since 1890, though many were never released. Published and unpublished records show
that 154 were released, almost all of them (151) as biological control agents of insect
pests and weeds. An estimated 24.5% and 66.7% of the species released against insect
pests and weeds, respectively, established populations in Florida. The proportion of
insect predators (26.7%) was very similar to that of insect parasitoids (23.9%) estab-
lished. Insect pests targeted were mainly Homoptera (48%), Lepidoptera (24%), and
Coleoptera (10%). Most of the insect pests (79%) and weeds (75%) targeted are not
native to Florida; 43% of the insect pests are native to Asia, and 50% of the weeds are
native to South America. None of the native insect pests and weeds targeted occurs
only in Florida. There was no clear relationship of the number of individuals released,
nor of their geographic origin, nor of the county in which they were released, to the
probability of establishment.

RESUME

Desde 1890, cerca de 351 species de insects han sido introducidas en Florida con
la intenci6n de ser liberadas, aunque muchas de ellas nunca lo fueron. Reports pub-
licados, asi como ineditos, muestran que 154 species fueron liberadas en el campo,
casi todas (151) como agents de control biol6gico contra insects plagas y malezas. Se
calcula que un 24,5% de las species liberadas contra insects plaga y un 66,7% de las
species liberadas contra malezas lograron establecer poblaciones en Florida. La prop-
orci6n de species depredadoras establecidas (26,7%) es muy similar a la proporci6n
de species parasitoides estableGidas (23,9%). Las plagas contra las cuales las
liberaciones fueron hechas, son principalmente Homoptera (48%), Lepidoptera (24%)
y Coleoptera (10%). La mayoria de los insects plaga (79%) y malezas (75%) involuc-
radas no son nativas de Florida; 43% de los insects plaga son originarios de Asia y 50
de las malezas son nativas de Sur Am6rica. Ninguna de estas species de insects plaga
o malezas ocurren exclusivamente en Florida. No existe una clara relaci6n entire la
probabilidad de establecimiento del agent de control biol6gico y el n6mero de indi-
viduos liberados, su origen geografico o el condado donde se liberaron.



Insects have been introduced into countries around the world for various purposes.
Perhaps the most common purpose, at least in this century, has been for biological
control (see Sailer 1972, van den Bosch & Messenger 1973, DeBach 1974, DeBach &










Florida Entomologist 76(1)


Rosen 1976). Perkins & Swezey (1924) and Imms (1926) provide insights into.some of
the earliest organized biological control efforts in the USA, in Hawaii.
Documenting the history of insect introductions into any region is a daunting, yet
important task. Laing & Hamai (1976), Clausen (1978), Luck (1981), and Funasaki et
al. (1988) provide examples of tabulations of the biological control agents introduced
into various regions. Denmark (1964) and Denmark & Porter (1973) previously have
documented the introduction of biological control agents into Florida, and here we
expand the scope of documentation, and bring it up-to-date. The current review comple-
ments our previous review of Florida's recent immigrant insects (Frank & McCoy 1992).

METHODS

We chose to include in our list of introduced insects all taxa that could be verified
as legitimate introductions (see Frank & McCoy 1990). Where some question exists as
to the legitimacy of a particular introduction, we have so indicated. Because most intro-
duced insects were imported as biological control agents, we also have included a list
of targets of these agents. It should not be assumed that the species included on the
list of targets are necessarily important pests, nor that species not included on the list
are necessarily unimportant pests. Particularly in what might be considered the "early
days" of biological control, before World War II, insect species might have been consid-
ered important pests based on wrongful information [see Eichmann 1943, see Ignelater
luminosus (Illiger) (Coleoptera: Elateridae) in Table 2] or important pests might even
have been misidentified (see Clausen 1942, see Metaphycus helvolus (Compere)
(Hymenoptera: Encyrtidae) in Table 2].
We constructed our tabulations of introduced insects and of targets from both pub-
lished and unpublished records. We then verified the tabulation by consulting au-
thorities on the included taxa and biological control practitioners. We cannot guarantee
completeness, however. More specific information on the methods employed to con-
struct tabulations of the two kinds of organisms follows.

Targets of Classical Biological Control Efforts

Our tabulation of pest organisms that have been targeted in Florida for classical
biological control (Table 1) includes a variety of information. We have given, when we
have the information, the pest organisms' most widely used common names, their prob-
able geographical origins, their exact (when documented) or probable decades of arrival
in Florida, and the economic and/or ecological beneficiaries of their control (e.g., crops
attacked).
Some biological control agents were imported with the notion of releasing them
against Parlatoria ziziphi (Lucas) and Toxoptera citricida (Kirkaldy). Any releases of
these agents ultimately were made against other pests, however (T. citricida does not
even occur in Florida), so we have not included them in our tabulation. Other agents
were introduced against Heliothis spp. and Spodoptera spp., but without specification
of which particular species were the intended targets. We refrained from guessing, and
included in our tabulation only the species of those genera that we know were targets.
We tried to distinguish between adventive pests and those that are native to Florida,
although a few ambiguities remain. Adventive pests are those of foreign origin, and
they may be subdivided into those that immigrated and those that were introduced (see
Frank & McCoy 1990 for definitions of these and related terms).
For some immigrant pests, a record of their arrival in Florida exists. For example,
Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) migrated eastward from
the Rocky Mountains after the widespread planting of potatoes provided a food source.


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy


TABLE 1.PEST ORGANISMS THAT HAVE BEEN TARGETED IN FLORIDA FOR CLASS-
ICAL BIOLOGICAL CONTROL, WITH THEIR COMMON NAMES, PROBABLE
ORIGINS, EXACT OR PROBABLE DECADE OF ARRIVAL INFLORIDA, AND
ECONOMIC/ECOLOGICAL IMPACTS. A = ADVENTIVE, N = NATIVE, NA =
NORTH AMERICA, CA = CENTRAL AMERICA, SA = SOUTH AMERICA,
AF = AFRICA, As = ASIA, Au = AUSTRALIA, Eu= EUROPE, WI =
WEST INDIES, FIN = FREQUENT IMMIGRANT FROM THE NORTHERN
USA (EUROPEAN ORIGIN), FIS = FREQUENT IMMIGRANT FROM THE WEST
INDIES AND/OR OVERWINTERS IN SOUTH FLORIDA.

COLEOPTERA: CHRYSOMELIDAE
Leptinotarsa decemlineata (Say), Colorado potato beetle. A: NA, 1920, potato
COLEOPTERA: COCCINELLIDAE
Epilachna varivestis Mulsant, Mexican bean beetle. A: NA/CA, 1920, beans
COLEOPTERA: CURCULIONIDAE
Anthonomus grandis Boheman, boll weevil. A: NA, 1910, cotton
Asynonychus godmanni Crotch, Fuller rose weevil. A: SA, 1870, citrus/ornamentals
Diaprepes abbreviatus (L.), Apopka weevil OR cane root borer. A: WI, 1960, citrus/
sugarcane
Hypera postica (Gyllenhal), alfalfa weevil. A: Eu, 1970, alfalfa and other legumes
COLEOPTERA: SCARABAEIDAE
Euetheola humilis (LeConte), sugarcane beetle. N, sugarcane
DIPTERA: CULICIDAE
Aedes aegypti (L.), yellow fever mosquito. A: Af, <1850, humans
DIPTERA: MUSCIDAE
Musca domestic L., house fly. A: Af, <1850, humans/livestock
Stomoxys calcitrans (L.), stable fly. A: Af, 1900, livestock
DIPTERA: TEPHRITIDAE
Anastrepha suspense (Loew), Caribbean fruit fly. A: WI, 1920, fruits/citrus
HEMIPTERA: PENTATOMIDAE
Nezara viridula (L.), southern green stinkbug. A: Af, <1850, vegetables
HEMIPTERA: TINGIDAE
Leptodictya tabida (Herrich-Schaeffer), sugarcane lace bug. A: NA/CA/SA, 1990, sugar-
cane
HOMOPTERA: ALEYRODIDAE
Aleurocanthus woglumi Ashby, citrus blackfly. A: As, 1930/1970, citrus
Aleurodicus disperus Russell, spiralling whitefly. A: WI/CA/SA, 1950, citrus/coconut
Bemisia tabaci Gennadius, sweetpotato whitefly. A: As, 1890, field crops/ornamentals
Dialeurodes citri (Ashmead), citrus whitefly. A: As, 1870, citrus/ornamentals
Dialeurodes citrifolii (Morgan), cloudywinged whitefly. A: As, 1900, citrus
HOMOPTERA: APHIDIDAE
Acyrthosiphon pisum (Harris), pea aphid. A: Eu/As, 1900, peas/alfalfa
Aphis gossypii Glover, cotton aphid. N, cotton/curcurbits
Aphis spiraecola Patch, spirea aphid. A: As, 1920, citrus
Myzus persicae (Sulzer), green peach aphid. N, fruits
Sipha flava (Forbes), yellow sugarcane aphid. N, sugarcane
Therioaphis maculata (Buckton), spotted alfalfa aphid. A: Eu/As, 1960, alfalfa/clover
Toxoptera aurantii (Fonscolombe), black citrus aphid. ?, ?, citrus









Florida Entomologist 76(1)


TABLE 1. (Continued)

HOMOPTERA: COCCIDAE
Ceroplastes cirripediformis Comstock, barnacle scale. A: As, 1870, citrus/ornamentals
Coccus hesperidum L., brown soft scale. A: As, 1870, citrus
Saissetia neglecta DeLotto, Caribbean black scale. N?, citrus
HOMOPTERA: DELPHACIDAE
Perkinsiella saccharicida Kirkaldy, sugarcane leafhopper. A: Au,1980, sugarcane
Saccharosydne saccharivora (Westwood), West Indian canefly. A: WI, ?, sugarcane

HOMOPTERA: DIASPIDIDAE
Aonidiella aurantii (Maskell), California red scale. A: As, 1890, citrus
Aspidiotus destructor Signoret, coconut scale. A: As, 1910, coconut
Chrysomphalus aonidum (L.), Florida red scale. A: As, 1880, citrus
Fiorinia theae Green, tea scale. A: As, 1910, camellia/citrus
Lepidosaphes beckii (Newman), purple scale. A: As, 1850, citrus
Pseudaulacaspis cockerelli (Cooley), false oleander scale. A: As, 1940, ornamentals
Pseudaulacaspis pentagon (Targioni-Tozzetti), white peach scale. A: As, 1880, fruits/
ornamentals
Unaspis citri (Comstock), citrus snow scale. A: As, 1880, citrus
Unaspis euonymi (Comstock), euonymus scale. A: As, 1960, ornamentals
HOMOPTERA: MARGARODIDAE
Icerya purchase Maskell, cottony cushion scale. A: Au, 1890, citrus
HOMOPTERA: PSEUDOCOCCIDAE
Antonina graminis Maskell, Rhodesgrass mealybug. A: As, 1940, grasses
Dysmicoccus boninsis (Kuwana), gray sugarcane mealybug. A: As, 1960, sugarcane
Dysmicoccus brevipes (Cockerell), pineapple mealybug. A: SA, 1910, sugarcane/pine-
apple
Planococcus citri (Risso), citrus mealybug. A: As, 1890, citrus
Saccharicoccus sacchari (Cockerell), pink sugarcane mealybug. A: Af, 1940, sugarcane
HYMENOPTERA: DIPRIONIDAE
Neodiprion lecontei (Fitch), redheaded pine sawfly. N, pine trees
HYMENOPTERA: FORMICIDAE
Solenopsis invicta Buren, red imported fire ant. A: SA, 1940, humans
LEPIDOPTERA: GELECHIIDAE
Pectinophora gossypiella (Saunders), pink bollworm. A: As, 1950, cotton
LEPIDOPTERA: GEOMETRIDAE
Epimecis detexta (Walker), avocado looper. N, avocado
LEPIDOPTERA: LASIOCAMPIDAE
Malacosoma disstria (F.), forest tent caterpillar. N, trees
LEPIDOPTERA: LYMANTRIIDAE
Lymantria dispar (L.), gypsy moth. A: FIN, trees
LEPIDOPTERA: NOCTUIDAE
Alabama argillacea (Hubner), cotton leafworm. A: SA/FIS, cotton
Anticarsia gemmatalis HGbner, velvetbean caterpillar. A: FIS, field crops
Helicoverpa zea (Boddie), corn earworm. N, cotton/field crops
Heliothis virescens (Boddie), tobacco budworm. N, field crops
Pseudoplusia includes (Walker), soybean looper. A: FIS, field crops


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy


Spodopterafrugiperda (J. E. Smith), fall armyworm. N, field crops
Trichoplusia ni Hiibner, cabbage looper. N, field crops
LEPIDOPTERA: PYRALIDAE
Diaphania hyalinata (L.), melonworm. A: FIS, cucurbits
Diaphania nitidalis Stoll, pickleworm. A: FIS, cucurbits
Diatraea saccharalis (F.), sugarcane borer. A: WI/CA/SA, 1860, sugarcane
Elasmopalpus lignosellus (Zeller), lesser cornstalk borer. N, field crops
EPIDOPTERA: YPONOMEUTIDAE
Plutella xylostella (L.), diamondback moth. A: Eu/Af, ?, cole crops
ORTHOPTERA: GRYLLOTALPIDAE
Scapteriscus abbreviatus Scudder, short-winged mole cricket. A: SA, 1900, grasses
Scapteriscus borellii Giglio-Tos, southern mole cricket. A: SA, 1900, grasses
Scapteriscus vicinus Scudder, tawny mole cricket. A: SA, 1900, grasses
THYSANOPTERA: THRIPIDAE
Selenothrips rubrocinctus (Giard), red-banded thrips. A: As, ?, fruits
ARALES: ARACEAE
Pistia stratiotes L., waterlettuce. N?, waterways
CAROPHYLLALES: AMARANTHACEAE
Alternanthera philoxeroides (Martius) Grisebach, alligatorweed. A: SA, 1890, water-
ways
HALORAGALES: HALORAGACEAE
Myriophyllum spicatum L., Eurasian watermilfoil. A: Eu/As, 1960, waterways
HYDROCHARITALES: HYDROCHARITACEAE
Hydrilla verticillata (Lf.) Royle, hydrilla. A: Eu/As, 1960, waterways
LAMIALES: VERBENACEAE
Lantana camera L., lantana. N, pastures
LILLIALES: PONTEDERIACEAE
Eichhornia crassipes (Martius) Solms, waterhyacinth. A: SA, 1880, waterways
MYRTALES: MYRTACEAE
Melaleuca quinquenervia (Cavanilles) S.T. Blake, melaleuca. A: Au, 1900, wetlands
SAPINDALES: ANACARDIACEAE
Schinus terebinthifolius Raddi, Brazilian peppertree. A: SA, 1890, woodlands



Although it is native to the USA, it is not native to Florida. For another example,
Scapteriscus spp. (Orthoptera: Gryllotalpidae) arrived as immigrants in the ballast of
ships from southern South America. They are native neither to the USA nor to Florida.
For introduced pests, a written record of their arrival almost always exists. They
were brought to Florida for various reasons and either were liberated in the wild or
escaped into the wild. That they have become pests shows in retrospect that the reasons
for their introductions were ill-considered. Only one insect species occurs among these
introduced pests: Icerya purchase Maskell (Homoptera: Margarodidae). In this incident,
a citrus grower, unfamiliar with the concept of host-specificity, and before state regula-
tions prohibited such action, imported in 1893 a specialist biological control agent
[Rodolia cardinalis (Mulsant) (Coleoptera: Coccinellidae)], even though its prey, I.
purchase, did not then occur in Florida. The prey was imported simultaneously as food
for the biological control agent and, while the agent dispersed and perished, the prey









Florida Entomologist 76(1)


TABLE 2. INSECTS IMPORTED INTO FLORIDA. THREE SYMBOLS ARE USED TO INDI-
CATE THE FATE OF BIOCONTROL AGENTS THAT WERE RELEASED: =
SPECIES THAT BECAME ESTABLISHED, = SPECIES THAT DID NOT BE-
COME ESTABLISHED, AND ? = SPECIES RELEASED VERY RECENTLY
WHOSE FATE IS YET UNKNOWN. THIS CLASSIFICATION IGNORES RE-
LEASES OF INSECTS THAT COULD NOT BE DISTINGUISHED FROM SPECIES
ALREADY PRESENT IN FLORIDA, AND IT ASSUMES THAT INSECTS RE-
LEASED SEVERAL YEARS AGO, BUT THAT HAVE NOT BEEN REPORTED
SUBSEQUENTLY FROM FLORIDA, ARE NOT ESTABLISHED.

BLATTARIA: OXYHYDROIDAE
Gromphadorina sp., from Madagascar, brought to Florida without authorization by the
pet trade and sold to the public; four of these giant hissing roaches were released
in Tampa by a disgruntled owner who feared they would be confiscated and killed
by Florida Dept. of Agriculture & Consumer Services inspectors (Chen 1989). ?
COLEOPTERA: BRUCHIDAE
Lithraeus atronotatus (Pic), from Brazil, imported in 1989 by D. H. Habeck against
Schinus terebinthifolius, was not cultured and was not released (AD 1989, D. H.
Habeck pers. comm.).
COLEOPTERA: CARABIDAE
Calosoma argentinense Csiki, from Argentina, released in 1941 and 1943-1944 and 1945-
1946 against Spodopterafrugiperda, not established (Annand 1945, 1947, Clausen
1956, Gross & Pair 1986). ~
Calosoma sycophanta L., from Europe via the northeastern USA, 16 were released in
1915 in Alachua County, against Anticarsia gemmatalis, not established (Watson
1917, Dowden 1962). ~
Pheropsophus aequinoctialis (L.), from Uruguay, Brazil and Bolivia, imported into
quarantine in 1986-1989, reared in quarantine and studied as a potential natural
enemy of Scapteriscus spp. (Gryllotalpidae), but the studies were not completed
due to lack of funds and are unpublished, and the surviving specimens ultimately
were killed in quarantine (Frank 1990).
Scarites sp., from Trinidad, misidentified by shipper as Pheropsophus and imported in
1987, terminated in quarantine (AD 1987).
Stenaptinus jessoensis (Morawitz), from Japan, a known natural enemy of Gryllotalpa,
imported in 1986 by J. H. Frank and studied in quarantine in comparison with
Pheropsophus aequinoctialis; the specimens were reared through several gener-
ations on eggs of Scapteriscus, but ultimately all died in quarantine (Frank 1990).
COLEOPTERA: CHRYSOMELIDAE
Agasicles hygrophila Selman & Vogt, from Argentina, released in 1965-1972 against
Alternanthera philoxeroides, in Alachua, Baker, Bradford, Broward, Calhoun,
Citrus, Clay, Dixie, Duval, Escambia, Flagler, Glades, Hendry, Hillsborough,
Marion, Palm Beach, Polk, Putnam, St. John's, St. Lucie, and Volusia counties,
established; from Argentina via California, imported in 1974 by N. R. Spencer;
from Argentina, imported and released in 1979 in Alachua County by G. R.
Buckingham (Zeiger 1967, Denmark & Porter 1973, AD 1974, Coulson 1977,
Buckingham et al. 1983, Buckingham & Habeck 1990). *
Octotoma scabripennis Gu6rin, from the Neotropical region via Australia, imported
into quarantine in 1981-1982 as a potential biocontrol agent for Lantana camera,
but died in quarantine (AD 1981, 1982, Habeck et al. in press, D. H. Habeck
pers. comm.).


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy


Uroplata girardi Pic, from the Neotropical region via Australia, imported into quaran-
tine in 1981 as a potential biocontrol agent for Lantana camera, but died in
quarantine (AD 1981, Habeck et al. in press, D. H. Habeck pers. comm.).
COLEOPTERA: CICINDELIDAE
Megacephala fulgida Klug, from Brazil, imported in 1985 by R. I. Sailer and studied
in quarantine as a potential natural enemy of Scapteriscus mole crickets, reared
through several generations by B. Munir and then by R. C. Hemenway, but
never released; the surviving specimens ultimately were killed (Frank 1990).
COLEOPTERA: COCCINELLIDAE
Brumoides suturalis (F.), from Pakistan via New Jersey, released in 1954-1955 against
Dialeurodes citri, aphids and other Homoptera on citrus and sugarcane in
Orange, Lake, Marion, Indian River, St. Lucie, and other counties, not estab-
lished (Selhime 1956, Denmark 1964, Charpentier et al. 1972, Gordon 1985,
Browning 1990). ~
Catana parcesetosa (Sicard), from Pakistan, released in 1956 from Orlando (Orange
County) against Dialeurodes citri, not established (Denmark 1964, Gordon 1985).

Chilocorus cacti (L.), from Jamaica, imported in 1989 by F. D. Bennett for laboratory
comparison with native material of this species, not released (AD 1989).
Chilocorus kuwanae Silvestri, from Japan via Maryland and Delaware, a predator of
Diaspididae, brought to Florida in 1989, by F. D. Bennett as a potential biocon-
trol agent of Fiorinia theae, Unaspis euonymi and U. citri, but material died in
quarantine (BIRL 1992, F. D. Bennett pers. comm.)
Chilocorus nigritus F., from Pakistan or India, imported in 1910 or 1911 by R. S.
Woglum against Aonidiella aurantii, but without record of release (Woglum
1913).
Coccidophilus sp., perhaps citricola Brethes, from Brazil, imported in 1989 by H. W.
Browning against Unaspis citri, but adults died in quarantine (AD 1989, H. W.
Browning pers. comm.).
Coccinella septempunctata L., from India, released in Gadsden County in 1958 against
Myzus persicae and Therioaphis maculata, not established; 3,400 eggs and larvae
released in 1959 in two fields against Sipha flava, not established, even though
it fed on this prey in the laboratory; approved for importation in 1960 under the
name "C. punctata", and approved for importation in 1961; later established in
the northeastern USA and brought from New Jersey via Delaware to Florida by
R. I. Sailer in 1976 and released in 1976-1977 against Acyrthosiphon pisum, now
established and spread widely in Florida (Denmark 1964, Charpentier et al. 1972,
Denmark & Porter 1973, Angalet & Jacques 1975, Angalet et al. 1979, Gordon
1985, Mizell & Tedders 1990, BIRL 1992). *
Coelophora inaequalis (F.), from Hawaii via Puerto Rico, 19 adults were released at
two sites against Sipha flava in 1939, establishment was uncertain at first, but
was clear by the late 1970s (Charpentier et al. 1972, Clausen 1956, 1978, Gordon
1985, Bennett et al. 1990). *
Cryptognatha gemellata Mulsant, from Trinidad, released in 1936 against Aspidiotus
destructor, not established (Dohanian 1937, Gordon 1985). -
Cryptognatha nodiceps Marshall, from Trinidad, released in 1936 and 1938 against As-
pidiotus destructor in Dade County, established (Dohanian 1937, Bartlett 1938,
Clausen 1956, Rosen & DeBach 1978, Gordon 1985). *
Cryptolaemus montrouzieri Mulsant, from Australia via California, imported in 1930
against Planococcus citri, established (Watson & Thompson 1933, Muma 1955,
Denmark 1964, Gordon 1985, Browning 1990). *









Florida Entomologist 76(1)


TABLE 2. (Continued)

Delphastus catalinae (Horn), from California, 12 individuals imported in September
1916, reared in a laboratory and released into an insectary; more were imported
in 1917 and released against Dialeurodes citri; the finding of numerous Delphas-
tus individuals in 1918 at a release site prompted distribution to other sites in
several counties (Watson 1917, 1918a,b, 1919, 1920, 1921, 1922, 1923a, 1924,
Merrill 1922, Muma 1955, Denmark 1964, Browning 1990). The identity of this
insect is an enigma because D. catalinae is now restricted to a small area of
California, while a second species [D. pallidus (LeConte)] was described from
Florida in 1878 and is restricted to Florida, and a third [D. pusillus (LeConte)]
occurs throughout the southern tier of states (Gordon 1985). No voucher material
of the insects imported from California seems to exist; no specimens of D.
catalinae could be found in the Florida State Collection of Arthropods in 1992,
though specimens collected at Cortez, Manatee County, on 14-1-1918 are in the
Florida State Collection of Arthropods. The last were identified by H. B.
Swartsel (the collector) as D. catalinae, but in 1921 were identified by A. J.
Mutchler (American Mus. Nat. Hist.) as D. pusillus. A letter from Mutchler to
Merrill concerning them is in the archives of the Florida Department of Agricul-
ture and Consumer Services. The explanation may be that Watson and Merrill
deluded themselves into believing that D. catalinae had become established in
Florida but they were dealing with specimens of D. pusillus, which is native
to Florida. ~
Delphastus pusillus (LeConte), from California, imported by J. R. Watson in 1917
under its junior synonym D. sonoricus Casey against Dialeurodes citri, though
not specified whether released, but D. pusillus is a widespread species which
already was present in Florida (Merrill 1922).
Delphastus sp. A. ("mottled brown"), from Puerto Rico, imported by H. Spencer (Sub-
tropical Insect Laboratory, USDA, Orlando) in May 1938 against Aspidiotus
destructor, 800 individuals, 738 were released, not established (Bartlett 1938,
1939, Clausen 1956, Rosen & DeBach 1978); this may have been Delphastus
nebulosus Chapin (F. D. Bennett, pers. comm.). ~
Delphastus sp. B. ("black"), from Puerto Rico, imported by H. Spencer (Subtropical
Insect Laboratory, USDA, Orlando) in May 1938 against Aspidiotus destructor,
49 individuals, 44 were released, not established (Bartlett 1938, 1939, Clausen
1956, Rosen & DeBach 1978; this may have been Zilus variipennis (Sicard)
and/or Z. gilvifrons Chapin (F. D. Bennett, pers. comm.). -
Egius platycephalus Mulsant, from Puerto Rico, imported in 1988 by F. D. Bennett
against Parlatoria ziziphi, failed to breed, and no releases were made (AD 1988).
Harmonia dimidiata (F.), from China via California, released in 1925-1926 against
Aphis spiraecola, established [mentioned in earlier literature at first as Leis
conformis (Boisduval), presumably by confusion of identity, and then as Leis
dimidiata 15-spilota and L. d. quinquedecimmaculata (Hope)], established (Wat-
son & Thompson 1933, 1940, 1941, Denmark 1964, Clausen 1978, Gordon 1985,
Browning 1990). *
Hippodamia convergens Gu6rin, perhaps from Texas, released in 1930 against aphids;
permit issued in 1966 for its importation and release against aphids, but is native
to Florida (Denmark 1964, Denmark & Porter 1973 (F. D. Bennett, pers. comm).
Hippodamia variegata (Goeze), from India, 900 adults and 7,950 eggs were released in
Gadsden and Palm Beach counties and perhaps other localities in 1957 against
Acyrthosiphon pisum, Myzus persicae, Siphaflava, Therioaphis maculata, and
other aphids, 94 adults and 212 eggs in 1958, not established (Denmark 1964,
Charpentier et al. 1972, Gordon 1985, Jackson 1990). ~


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy 9

Menochilus sexmaculatus (F.), from Pakistan or India, imported in 1910 or 1911 against
[citrus] aphids by R. S. Woglum, released; from India via Texas, imported in
1954 by A. G. Selhime, released in 1955 in Indian River, St. Lucie, Lake, Marion,
Orange and Seminole counties against "citrus aphids", not established (Essig
1931, Selhime 1955, Gordon 1985). ~
Nephaspis oculata (Blatchley), from Hawaii, 600 released in 1982 in Broward County
and 600 in Dade County under the erroneous name Nephaspis amnicola by R.
I. Sailer and C. R. Thompson against Aleurodicus dispersus, but occurred al-
ready in Florida (AD 1982, ROBO 1982, Bennett & Noyes 1989).
Nephus binaevatus (Mulsant), from California, imported in 1923 by J. R. Watson against
mealybugs, but not specified whether released, and not now present in Florida
(Watson 1923b).
Pentilia castanea Mulsant, from Trinidad, over 600 individuals were imported and the
survivors were released in 1936 against Aspidiotus destructor, but the species
did not become established (Dohanian 1937); there is uncertainty over the identifi-
cation because specimens were identified as above by USDA taxonomists,
whereas in Trinidad the species was recorded as P. insidiosa Mulsant (Dohanian
1937); a species under the latter name from Trinidad was distributed to Bermuda,
Fiji, Principe, and Puerto Rico (Bartlett 1978). ~
Pentilia egena Mulsant, from Brazil, imported in 1989 by H. W. Browning against
Unaspis citri, but no releases were made (AD 1989, H. W. Browning pers.
comm.).
Pseudoazya trinitatis (Marshall), from Trinidad via Puerto Rico, released in 1938
against Aspidiotus destructor in Dade County; specimens were found in 1939 but
not subsequently (Bartlett 1938, Clausen 1956, Rosen & DeBach 1978, Gordon
1985). ~
Rhyzobius lophanthae (Blaisdell), perhaps from Australia via California, but no records
of the introduction into Florida have been located; this is an Old World species
which either was introduced into Florida or immigrated from California (Gordon
1985).
Rodolia cardinalis (Mulsant), from Australia via California, released in 1893 by a com-
mercial plant-growing business in Pinellas County, in misunderstanding of its use
in biological control, but together (inadvertently) with its prey, Icerya purchase,
which resulted not in its establishment, but in establishment of its prey, causing
a severe infestation, the first in Florida (Berger 1915); after ill-conceived at-
tempts to control the pest by physical methods, R. cardinalis was reintroduced
in 1899 from California and established in Pinellas County, leading to highly
successful biological control. There surely cannot be any more perfect example
of the principles of biological control. This example provides primafacie evidence
of the need for regulation of importation of biocontrol agents; a further importa-
tion and release was approved in 1971 (Berger 1915, Merrill 1922, Denmark 1964,
Denmark & Porter 1973, Mizell 1990). *
Scymnus nubilis Mulsant, from India, released (from Quincy) in Gadsden County in
1957 against Myzus persicae and Therioaphis maculata; in 1957, 66 adults and
126 eggs (from India) were released at at one site, and in 1958, 3,200 eggs were
released at three sites, and in 1959, 800 adults were released at one site from
Belle Glade (Palm Beach County) against Sipha flava, not established, even
though the species was reared successfully in the laboratory on Sipha flava
(Clausen 1959, Denmark 1964, Charpentier et al. 1972, Gordon 1985, Jackson
1990). ~
Serangium flavescens (Motschulsky) from Pakistan, imported in 1910 against
Dialeurodes citri by R. S. Woglum, but specimens were dead upon arrival, im-
ported in 191 by R. S. Woglum, but arrived in winter when prey was not avail-









Florida Entomologist 76(1)


TABLE 2. (Continued)

able, and all died before releases could be made (Woglum 1913, Gordon 1985,
Browning 1990).
Stethorus utilis (Horn) (as S. atomus Casey), origin not stated, released in 1956 from
Weirsdale (Marion County), but is native to Florida (Denmark 1964).
Sticholotis madagassa Weise, from India, an unsolicited shipment received in 1979,
presumably against Pseudaulacaspis spp., dead on arrival in Florida (AD 1979).
Zagloba aeneipennis (Sicard), from Trinidad, released in 1936 against Aspidiotus de-
structor, not established (Dohanian 1937, Gordon 1985). -
COLEOPTERA: CURCULIONIDAE
Bagous affinis Hustache, from India, imported in 1982-1983, 1986, and 1990-1991 by C.
A. Bennett and G. R. Buckingham against Hydrilla verticillata, released in 1987
in Osceola County, established temporarily (AD 1982, 1983, 1986, 1990, 1991,
Buckingham 1988, Buckingham & Habeck 1990). *
Bagous dilgiri Vazirani, from India, imported in 1983 by G. R. Buckingham against
Hydrilla verticillata, terminated in quarantine (AD 1983, Bennett & Buckingham
1991, G. R. Buckingham pers. comm.).
Bagous laevigatus O'Brien & Pajni, from India, imported in 1983 and 1986 by G. R.
Buckingham against Hydrilla verticillata, terminated in quarantine after host-
range testing (AD 1983, 1986, G. R. Buckingham pers. comm.).
Bagous vicinus Hustache, from India, imported in 1983 by G. R. Buckingham against
Hydrilla verticillata, terminated in quarantine (AD 1983, Bennett & Buckingham
1982, G. R. Buckingham pers. comm.).
Bagous n. sp., from Australia, imported in 1987, 1988, and 1991 by G. R. Buckingham
against Hydrilla verticillata, released in 1991 in Broward, Palm Beach and Sum-
ter counties, establishment still uncertain (AD 1987, 1988, 1991, Center 1992, G.
R. Buckingham pers. comm.). ?
Eubrychius sp., from China, imported in 1991 by G. R. Buckingham against Myriophyl-
lum spicatum, but culture was lost in quarantine (AD 1991, G. R. Buckingham
pers. comm.).
Neochetina bruchi Hustache, from Argentina, imported in 1974 by N. R. Spencer
against Eichhornia crassipes, released in 1974, established; imported in 1975 by
G. E. Allen and released in Lee County (AD 1974, 1975, Perkins & Maddox 1976,
Grissell 1978, Cassani et al. 1981, Center & Durden 1986, Haag 1986, Bucking-
ham & Habeck 1990). *
Neochetina eichhorniae Warner, from Argentina, released in 1972 first in Broward
County against Eichhornia crassipes, established; released in 1974 in Glades and
Lee counties; imported in 1975 by G. E. Allen (Perkins 1973, AD 1975, Cassani
et al. 1981, Center & Durden 1986, Haag 1986, Buckingham & Habeck 1990). *
Neohydronomus affinis Hustache, from Brazil via Australia, imported in 1986 and 1988
by D. H. Habeck against Pistia stratiotes, released in 1987-1988 in Broward,
Palm Beach, and St. Lucie counties, established [mentioned earlier by error as
N. pulchellus Hustache], now widely distributed in Florida (AD 1986, 1988,
Thompson & Habeck 1989, Buckingham & Habeck 1990, Dray et al. 1990, D. H.
Habeck pers. comm.). *
Omolabus piceus Germar, from Brazil, imported in 1989 by D. H. Habeck against
Schinus terebinthifolius, not released (AD 1989, D. H. Habeck pers. comm.).
Oxyops vitiosa Pascoe, from Australia, imported into quarantine in 1992 as a potential
biocontrol agent for Melaleuca quinquenervia, not released (Habeck et al. in
press, D. H. Habeck pers. comm.).
Phytobius leucogaster (Marsham), from California, imported in 1978-1979 by G. R.


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy 11

Buckingham against Myriophyllum spicatum, released in 1979 in Levy County,
probably not established (AD 1978, 1979, Buckingham & Habeck 1990, G. R.
Buckingham pers. comm.). ~
COLEOPTERA: ELATERIDAE
Ignelater luminosus (Illiger), from Puerto Rico, released in 1943 against scarab larvae,
especially Euetheola humilis [given as E. rugiceps (LeConte), which is a
synonym] attacking sugarcane, though it is other genera and species of
Scarabaeidae, not this one, which have been implicated as pests of sugarcane in
Florida; not established (Annand 1944, Clausen 1956).
COLEOPTERA: PSELAPHIDAE
Fustiger elegans Raffray, from Argentina, imported in 1987 by D. P. Wojcik against
Solenopsis invicta, not released (AD 1987, D. P. Wojcik pers. comm.).
DIPTERA: CECIDOMYIIDAE
genus and species indet. (three species), from Brazil, imported in 1992 by F. D. Bennett
as potential biocontrol agents for Schinus terebinthifolius (F. D. Bennett pers.
comm.).
DIPTERA: CHIRONOMIDAE
Polypedilum dewulfi Goetghebuer, from Burundi, imported in 1990 by G. R. Buckingam
& C. A. Bennett against Hydrilla verticillata (AD 1990, G. R. Buckingham pers.
comm).
Polypedilum wittae (Freeman), from Burundi, imported in 1990 by G. R. Buckingam
& C. A. Bennett against Hydrilla verticillata (AD 1990, G. R. Buckingham pers.
comm).
DIPTERA: CRYPTOCHETIDAE
Cryptochetum iceryae (Williston) [not C. monophlebi Skuse], from Australia via Califor-
nia, imported in 1917 and released against Icerya purchase in Pinellas County,
and reported as established (Thorpe 1930, Denmark 1964, Bartlett 1978), but not
seen in recent years (F. D. Bennett pers. comm.). *
DIPTERA: CULICIDAE
Toxorhynchites amboinensis (Doleschall), from Indo-Malaysia via Louisiana, released
ca. 1986 and in subsequent years in Duval and St. Lucie counties against Aedes
aegypti and other mosquitoes whose larvae inhabit artificial containers, does not
survive winters and is not established (G. A. Curtis pers. comm., E. Schreiber
pers. comm.). ~
Toxorhynchites splendens (Wiedemann), from Burma via Hawaii, Indiana, and
Louisiana, released in Bay County in 1986-1988, in Leon and Sarasota counties
in 1990, in Palm Beach and Walton counties in 1991, and in Hillsborough County
in 1992, against Aedes aegypti and other mosquitoes whose larvae inhabit artifi-
cial containers, has not survived the winter at any of the localities (E. Schreiber,
pers. comm.). -
DIPTERA: EPHYDRIDAE
Hydrellia balciunasi Bock, from Australia, imported in 1988, 1989 and 1991 by G. R.
Buckingham against Hydrilla verticillata, released in 1989 in Broward County,
in 1990 in Broward County, and in 1991 in Broward, Collier, and Sumter counties,
establishment not certain (AD 1988, 1989, 1991, Buckingham & Habeck 1990,
Center 1992, G. R. Buckingham pers. comm). ?
Hydrellia pakistanae Deonier, from India, Pakistan, and China, imported in 1986 and
1990 by G. R. Buckingham against Hydrilla verticillata, released in 1987 in Polk
and Marion counties, in 1988 in Broward, Glades, and Palm Beach counties, in









Florida Entomologist 76(1)


TABLE 2. (Continued)

1989 in Broward, Glades, Osceola, and Polk counties, in 1990 in Broward, Glades,
Jefferson, Lake, Okeechobee, Osceola, and Palm Beach counties, established
(AD 1986, 1990, Buckingham 1988, Buckingham & Habeck 1990, Center 1992, G.
R. Buckingham pers. comm.). *
Hydrellia sp., from India, China, and Japan, imported in 1990, 1991, and 1992 by G.
R. Buckingham against Hydrilla verticillata, not yet released (AD 1990, G. R.
Buckingham pers. comm.).
DIPTERA: SARCOPHAGIDAE
Heliocobia rapax (Walker), from Cuba, 65 puparia were imported in 1927 and released
against Diatraea saccharalis; there were no recoveries (Charpentier et al. 1972).

DIPTERA: TACHINIDAE
Archytas incertus (Macquart), from Argentina, released in 1944 against Spodoptera
frugiperda, not established (Clausen 1956, Gross & Pair 1986). ~
Archytas sp. (as Pseudoarchytopsis sp.), from Uruguay, imported in 1943-1944 against
Spodopterafrugiperda (Annand 1945).
Compsilura concinnata (Meigen), from Europe via northeastern USA, 896 shipped to
Alachua County in 1915, and 2,125 in 1916, against Spodoptera frugiperda, but
numbers (if any) released seem not to have been recorded, not established (Dow-
den 1962). -
Evbrissa vittata (Meigen) (as Phania vittata), from Japan via Delaware, imported in
1976 by W. H. Whitcomb as a potential biocontrol agent for Nezara viridula,
though not reported by Jones (1988) as a parasitoid of this host (BIRL 1992).
Hemisturmia sp., from Puerto Rico, imported in 1991 by J. L. Capinera and H. A.
Smith against Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera
pers. comm.).
Incamyia cuzcensis Townsend, from Paraguay and Peru, imported in 1979 against
noctuids (AD 1979).
Incamyia sp., from Paraguay and Peru, imported in 1979 against noctuids (AD 1979).
Lixophaga diatraeae (Townsend), from Cuba, Puerto Rico, and Trinidad, released in
1926-1927 (about 3,000 adults, from Cuba, at three sites) and 1936 (160 adults,
from Puerto Rico, at three sites, in Hendry, Indian River, and Palm Beach
counties) and 1938-1939 (27,292 adults, from Cuban stock, at one site) and 1948
(661 adults, from Cuba, at several sites) and 1949 (551 adults, from Cuba, at
several sites) and 1950-1961 (82,674 adults, from Cuban stock, on one plantation
in Indian River County: 11,441 in 1950, 10,185 in 1951, 8,703 in 1952, 9,803 in
1953, 11,555 in 1954, 10,914 in 1955, 11,937 in 1956, 8,786 in 1957, and 250 in
1961) and 1961 (491 puparia, from Trinidad, at one site) and 1967 (500 adults
reared from material collected in southern Louisiana whose origin was Trinidad,
in experimental plots) and 1969 (9,000 adults, from Trinidad, at several sites),
against Diatraea saccharalis, but no recoveries were made; from Barbados via
Trinidad, released in 1973 (28,500 adults in Palm Beach County and 18,000 in
Hendry County), and from Louisiana via Mississippi, in 1974 (60,000 in Palm
Beach County), augmentatively, but the program was discontinued; not recov-
ered in recent years (Watson 1928, Annand 1944, Gifford 1964, Charpentier et
al. 1967, 1972, Bennett 1971, Summers et al. 1976, Clausen & Oatman 1978, Hall
1986, Bennett et al. 1990, F. D. Bennett pers. comm.). ~
Lixophaga sphenophori (Villeneuve), from New Guinea (perhaps via Hawaii), released
in 1963 or 1964 against Diatraea saccharalis [(though it is a natural enemy of
Sphenophorus (Curculionidae)], not established (Gifford 1964). -


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy


Lydella thompsoni Herting, a palearctic species which is established in the northern
USA, released (erroneously under the name L. grisescens Robineau-Desvoidy)
in 1963 or 1964 against Diatraea saccharalis, not established (Gifford 1964). ~
Metagonistylum minense Townsend, from Brazil; 191 adults from stock imported from
Puerto Rico (origin Brazil) were released in 1938 at three sites, and in 1939-1942,
3,000 adults from Sao Paulo, Brazil were released (against Diatraea saccharalis
in Indian River County); recoveries were made in 1940-1941, 1943, and 1945, but
not subsequently, and not established (Holloway & Mathes 1942, Scaramuzza &
Ingram 1942, Annand 1944, Gifford 1964, Charpentier et al. 1972, Clausen 1978,
Bennett et al. 1990).-
Ormia depleta (Wiedemann), from Sao Paulo, Brazil, imported in 1985-1986 by R. I.
Sailer and in 1986-1989 by J. H. Frank, first bred by S. A. Wineriter in 1987
(Wineriter & Walker 1990), released in April 1988 in Alachua County and October
1988 in Manatee County against Scapteriscus spp., with subsequent releases in
1989-1992 in Baker, Broward, Citrus, Collier, Dade, Duval, Highlands, Hills-
borough, Lee, Marion, Okaloosa, Orange, Osceola, Palm Beach, Pasco, Pinellas,
Polk, St. John's, Sarasota, and Volusia counties, and now established widely in
peninsular Florida (Frank 1990, Parkman & Frank 1992). *
Paratheresia claripalpis (van der Wulp), from Peru, over 4,000 adults released in 1932
in Palm Beach and Indian River counties, 46 adults released in 1936 in Indian
River County, against Diatraea saccharalis, achieving up to 29% parasitism in
1936, but no longer present by 1964, when 390 adults from Trinidad were released
at one site, and 1969 when 750 adults from Trinidad were released at one site,
with only one specimen recovered; not established permanently (Jaynes 1938,
1939, 1939, Scaramuzza & Ingram 1942, Gifford 1964, Bennett 1971, Charpentier
et al. 1972, Clausen 1978, Bennett et al. 1990). -
Phorocera sp., from Argentina, imported in 1943-1944 against Spodoptera frugipe.'da
(Annand 1945).
Sturmiopsis inferens Townsend, from India via Delaware, 156 adults and puparia were
imported in 1962 by J. R. Gifford, 66 adults in 1964 by G. A. Mann, as a potential
biocontrol agent for Diatraea saccharalis, which was parasitized in the labora-
tory, but no releases were made; more were imported in 1971 by T. E. Summers;
a few were imported in 1974-1975 by N. R. Spencer among shipments of S.
parasitica, acting as quarantine clearance for shipments to the sugarcane zone
(Gifford 1964, Charpentier et al. 1972, AD 1974, 1975, BIRL 1992, N. R. Spencer
pers. comm.).
Sturmiopsis parasitica (Curran) from Ghana via India, imported in 1974-1975 by N. R.
Spencer, acting as quarantine clearance for shipments to the sugarcane zone,
against Diatraea saccharalis, intended for release (AD 1974, 1975, N. R. Spencer
pers. comm.).
Sturmiopsis sp., from India via Delaware, imported in 1964 by G. A. Mann as a biocon-
trol agent for Diatraea saccharalis (BIRL 1992).
Trichopoda pilipes (F.), from Hawaii (USA) via Delaware, imported in 1972 and 1973
by W. H. Whitcomb, from Montserrat via Delaware, imported in 1973 by W. H.
Whitcomb and N. R. Spencer as a biocontrol agent for Nezara viridula (AD 1973,
BIRL 1992, N. R. Spencer pers. comm.).
Trichopoda sp. (as Trichopodopsis argentinensis Blanchard), from Argentina via Dela-
ware, imported in 1974 by W. H. Whitcomb and N. R. Spencer as a potential
biocontrol agent for Nezara viridula, terminated in quarantine (AD 1974, BIRL
1992, N. R. Spencer pers. comm.).
Trichopoda sp., from Japan via Delaware, imported in 1976 by W. H. Whitcomb against
Nezara viridula (AD 1976).
genus and species indet., from Argentina via Delaware, imported in 1974 by N. R.










Florida Entomologist 76(1)


TABLE 2. (Continued)

Spencer against Alabama argillacea, terminated in quarantine (AD 1974, N. R.
Spencer pers. comm.).
genus and species indet., from Brazil, imported in 1975 by W. H. Whitcomb against
Anticarsia gemmatalis and Pseudoplusia includes (AD 1975).
genus and species indet., from Argentina, imported in 1979 against Spodoptera spp.
(AD 1979).
genus and species indet., from Colombia, emerged in quarantine from Urbanus proteus
L. which was imported by R. I. Sailer in 1982 as host for Ardalus sp. (AD 1982).
HEMIPTERA: MIRIDAE
Tytthus mundulus (Breddin), from Hawaii, imported in 1982-1984 by H. A. Denmark,
R. Nguyen, D. L. Harris, and O. Sosa against Perkinsiella saccharicida, re-
leased in 1982-1984 in Hendry and Palm Beach counties (201 in Palm Beach
County and 20 in Hendry County in 1982-1983), but not established (AD 1982,
1983, 1984, ROBO 1982, 1983, Nguyen et al. 1984, Bennett et al. 1990). ~
HEMIPTERA: PENTATOMIDAE
Cantheconidia furcellata (Wolff) (also as Eocanthecona furcellata (Wolff)), from Thai-
land, imported in 1980 by J.-M. Tseng and R. I. Sailer as a predator of
Malacosoma disstria, Anticarsia gemmatalis, Pseudoplusia includes
(Walker), and Leptinotarsa decemlineata (Say), released in 1981 in Alachua
(13,000) and Gadsden (756) counties (AD 1980, ROBO 1981, Stange 1982). -
HEMIPTERA: REDUVIIDAE
Zelus longipes (L.), from Jamaica via Trinidad, 275 adults were released in 1959 against
Saccharosydne saccharivora, but already was present in Florida (Bennett 1960,
Simmonds 1969, Charpentier et al. 1972, Bennett et al. 1990, F. D. Bennett pers.
comm.).
HYMENOPTERA: APHELINIDAE
Aphelinus asychis Walker (as A. semiflavus Howard), from Israel, 890 adults were
released in 1957 at one site against Sipha flava, and some specimens were ob-
tained later in the field, but without permanent establishment (Denmark 1964,
Charpentier et al. 1972). -
Aphelinus flavipes (Foerster), from Taiwan via Delaware, imported and released in
1972 and 1973 by N. R. Spencer as a biocontrol agent for Myzus persicae, re-
leased in Alachua County, but could not be differentiated from native species so
establishment uncertain (BIRL 1972, N. R. Spencer pers. comm.).
Aphelinus gossypii Timberlake, permit issued in 1969 for its importation and release
in Florida; from Hong Kong, imported as a biocontrol agent of Aphis spiraecola
in 1988 by F. D. Bennett, but lost in quarantine due to activity of an accompany-
ing hyperparasite, Tassonia gloriae Girault (Hym.: Encyrtidae), and to a short-
age of host material (Denmark & Porter 1973, AD 1988, Mizell & Tedders 1990).
Aphytis aonidiae (Mercet) (as A. citrinus Compere), permit issued in 1967 for its impor-
tation and release in Florida, probably against Aonidiella aurantii (Denmark &
Porter 1973).
Aphytis coheni DeBach, permit issued in 1967 for its importation into Florida, presum-
ably against Aonidiella aurantii (Denmark & Porter 1973).
Aphytis gordoni DeBach & Rosen, from Hong Kong via Delaware, imported in 1971 by
A. G. Selhime as a potential biocontrol agent for Unaspis citri (Selhime & Brooks
1979, BIRL 1992).
Aphytis holoxanthus DeBach, from Hong Kong via Israel, New Jersey, and California,
15,500 adults imported by D. W. Clancy in 1960 and released against Chrysom-
phalus aonidum in Lake, Orange, Polk, Hillsborough, and DeSoto counties, es-


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy


tablished by 1961 and very successful, and by mid-1964 had dispersed over the
entire citrus-growing area; imported in 1989 by H. W. Browning (as Aphytis sp.)
before specific identity was known, not released (Clancy et al. 1963, Denmark
1964, Selhime et al. 1969, Denmark & Porter 1973, Selhime & Brooks 1979,
McCoy 1985, Browning 1990, H. W. Browning pers. comm.). *
Aphytis lepidosaphes Compere, from China via California, released in 1958 against
Lepidosaphes beckii and Chrysomphalus aonidum, but had already arrived in
Florida as an immigrant, discovered in 1958, and become established (Clancy &
Muma 1959, Denmark 1964, Selhime & Brooks 1979, Rosen & DeBach 1978).
Aphytis lingnanensis Compere, from Hong Kong via New Jersey, imported in 1971 by
A. G. Selhime and C. W. McCoy, and in 1972 by A. G. Selhime, released in 1971
against Unaspis citri, believed to have become established, but inability to distin-
guish material of this species from that of a presumed native species (A. sp. nr.
lingnanensis) and three other imported strains of the same species from else-
where have made definitive conclusions impossible; one of these strains was a
hybrid between stock from Florida and from Puerto Rico, and the second was
collected from Aonidiella aurantii in California; the third of the strains, from
Thailand, was imported in 1989 (via Australia) and was released in 1990 by H.
W. Browning (AD 1989, Selhime & Brooks 1979, McCoy 1985, Browning 1990,
BIRL 1992, H. W. Browning pers. comm.).
Aphytis sp. nr. lingnanensis Compere, permit issued in 1972 for its importation and
release, presumably against Unaspis citri (Denmark & Porter 1973).
Aphytis melinus DeBach, permit issued in 1971 for its importation and release in
Florida, presumably against Aonidiella aurantii (Denmark & Porter 1973).
Aphytis theae (Cameron), from India, imported in 1975-1976 by N. R. Spencer against
Fiorinia theae, released in Alachua County in 1976-1978 by F. Collins and W.
H. Whitcomb, but did not survive the winters (AD 1975, 1976, Rosen-& DeBach
1977, Grissell 1978, Munir & Sailer 1985, Mizell 1990, Bennett & Capinera in
press). ~
Aphytis yanonensis DeBach & Rosen, from Japan via Texas, imported by F. D. Bennett
in 1987, released against Unaspis citri (AD 1987, F. D. Bennett pers. comm.). -
Aphytis sp., from China and Hong Kong, imported in 1986 by R. I. Sailer and in 1987
and 1988 by F. D. Bennett and J. H. Frank against Parlatoria ziziphi, died in
quarantine (AD 1986, 1987, 1988, F. D. Bennett pers. comm.).
Aphytis sp., from Egypt, imported by H. W. Browning against Parlatoria ziziphi, died
in quarantine (AD 1990, H. W. Browning pers. comm.).
Aphytis sp., from Hong Kong via Delaware, imported in 1973 by A. G. Selhime as a
potential biocontrol agent for Unaspis citri (Denmark & Porter 1973, BIRL
1992).
Aphytis sp., from India, imported in 1979 and 1980 by G. R. Buckingham against
Pseudaulacaspis cockerelli, not released (AD 1979, 1980, G. R. Buckingham
pers. comm., H. B. Glenn pers. comm.).
Aphytis sp., from India, unsolicited, in 1985, against Parlatoria ziziphi, could not be
cultured and was not released (AD 1986, H. B. Glenn pers. comm.).
Aphytis sp., from Japan via Texas, reared in quarantine on Aspidiotus nerii Bouch),
72,750 were released in Dade County against Unaspis citri, but not currently
distinguishable from existing species, so establishment uncertain (H. B. Glenn
pers. comm.).
Aphytis sp., from Puerto Rico, imported in 1990 against Parlatoria ziziphi, but not
reared and not released (AD 1990, F. D. Bennett pers. comm.).
Encarsia clypealis (Silvestri), from Mexico, imported in 1976 by A. G. Selhime, released
in Broward County against Aleurocanthus woglumi, not established (Dowell et
al. 1979, Browning 1990). ~










Florida Entomologist 76(1)


TABLE 2. (Continued)

Encarsiaformosa Gahan (most likely this species, from the neotropical region), brought
to Florida in 1928-1929 as a potential natural enemy of Dialeurodes citri, though
probably already occurred in Florida, where it attacks Trialeurodes and Bemisia
(Watson 1930, F. D. Bennett pers. comm.).
Encarsia sp. near formosa (Gahan), from Mexico, imported in 1990 by F. D. Bennett
against Bemisia tabaci, but not reared, and not released (AD 1990, F. D. Bennett
pers. comm.).
Encarsia sp. nr. haitiensis Dozier, from the Caribbean via Hawaii, imported in 1982
against Aleurodicus dispersus by R. I. Sailer and C. R. Thompson, 10 released
in 1982 in Broward County, but not established; also tested in quarantine against
Dialeurodes citrifolii (AD 1982, ROBO 1982, Stange 1986, Bennett & Noyes
1989). ~
Encarsia lahorensis (Howard), imported from Pakistan in the winter of 1911-1912 by
R. S. Woglum when host material was not available, and all died before releases
could be made (Woglum 1913); from Pakistan via California, imported in 1975 by
R. I. Sailer and placed in sleeve cages on trees in Alachua County; imported in
1977 by R. I. Sailer and released in Alachua County, later the same year in Polk
County, established, with subsequent dissemination to 64 more of the 67 counties
(e.g., released in Bay County in 1982 by J. A. Hogsette), against Dialeurodes
citri (AD 1975, Nguyen & Sailer 1979, Grissell 1979, ROBO 1982, Sailer et al.
1984, Nguyen 1986, Browning 1990). *
Encarsia lounsburyi (Berlese & Paoli), from India, released in 1977 against Fiorinia
theae, but was difficult to distinguish from a species already occurring in Florida
under the same name, and establishment is uncertain (Bennett & Capinera in
press, F. D. Bennett pers. comm.).
Encarsia lutea (Masi), from Israel, imported in 1990 by F. D. Bennett and R. Nguyen
against Bemisia tabaci, and from Sudan, imported in 1991 by F. D. Bennett (AD
1990, 1991).
Encarsia nigricephala Dozier, from Mexico, imported by F. D. Bennett in 1990 against
Bemisia tabaci, and from Guatemala, imported in 1991, but not bred and not
released (AD 1990, 1991, F. D. Bennett pers. comm.).
Encarsia opulenta (Silvestri), from India via Mexico (in part via Texas), by A. G.
Selhime, about 2,000 parasitoids were released in Broward County in 1976
against Aleurocanthus woglumi, established and highly successful (Hart et al.
1978, Grissell 1978, 1979, Selhime et al. 1982, McCoy 1985, Nguyen 1987, Brown-
ing 1990); released in Brevard (46,900), Lee (42,000), Martin (13,400) and
Okeechobee counties in 1981 by W. Grandberry (Dowell et al. 1979, ROBO 1981). *
Encarsia pergandiella Howard, from Mexico, imported in 1990 by F. D. Bennett (as E.
tabacivora) against Bemisia tabaci, and from Guatemala, imported in 1991, but
already was present in Florida, so no attempt was made to culture or to release
it (AD 1990, 1991, F. D. Bennett pers. comm.).
Encarsia sp. nr. protransvena Viggiani, from Puerto Rico, imported in 1988 by F. D.
Bennett against Dialeurodes citrifolii, but not released (AD 1988, F. D. Bennett
pers. comm.).
Encarsia sankarani Hayat, from India, imported into quarantine in 1976 by N. R.
Spencer as a potential biological control agent for Fiorinia theae; believed to
have been shipped to Dade County; there is no published record of its release,
but it now occurs in south Florida (AD 1976, Hayat 1989, Bennett & Capinera
in press, F. D. Bennett, pers. comm., N. R. Spencer pers. comm). *
Encarsia smith (Silvestri), from India via Mexico, imported in 1979 as a contaminant


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy


of E. clypealis cultures which in turn was imported as a biocontrol agent for
Aleurocanthus woglumi at a time when its behavior (female larvae are
parasitoids of Aleurocanthus, but male larvae are adelphoparasitoids of Encar-
sia) was not understood; released in 1979-1980 in several counties as a contami-
nant of laboratory cultures of Encarsia opulenta, in 1981 in Brevard (34) and Lee
(7) counties by W. Grandberry, established (ROBO 1981, Holder & McCluskie
1982, Nguyen et al. 1983, Browning 1990). *
Encarsia sp., permit issued in 1971 for its importation into Florida against an un-
specified pest (Denmark & Porter 1973).
Encarsia sp. (of Citrina Group) (as Aspidiotiphagus sp.), from Hong Kong via Dela-
ware, imported in 1972 by A. G. Selhime as a potential biocontrol agent for
Unaspis citri (BIRL 1992).
Encarsia spp., from the Cayman Islands, imported by F. D. Bennett in 1987 against
Dialeurodes citrifolii, and from Puerto Rico, imported by F. D. Bennett in 1987
and 1989, and from the Dominican Republic, imported in 1990 by H. W. Brown-
ing; was neither cultured nor released (AD 1987, 1989, 1990, H. W. Browning
pers. comm.).
Encarsia spp., from Brazil and Costa Rica and Grenada and Guadeloupe and Honduras
and Mexico and Puerto Rico, imported in 1990 by F. D. Bennett against Bemisia
tabaci, and from Brazil and Guatemala and Venezuela, imported in 1991 by F.
D. Bennett (AD 1990, 1991).
Encarsia sp., from Jamaica, imported in 1989 by F. D. Bennett and H. W. Browning
against Unaspis citri, not cultured and not released (AD 1989, H. W. Browning
pers. comm.).
Encarsia sp., from Puerto Rico, imported in 1987 and 1990 by F. D. Bennett against
Parlatoria ziziphi, not released (AD 1987, 1990).
Encarsia spp. (as Aspidiotiphagus sp.), from Hong Kong and China, imported in 1986,
1987, and 1988 by F. D. Bennett and J. H. Frank against Parlatoria ziziphi,
reared on Aspidiotus nerii, 1,400 released in 1987 in Dade County against
Chrysomphalus aonidum and Pseudaulacaspis cockerelli, 9,000 released in 1988
in Dade County against Pseudaulacaspis cockerelli and Unaspis citri, but not
currently distinguishable from existing species, so establishment uncertain; from
Hawaii in 1989, imported by F. D. Bennett, not released (AD 1986, 1987, 1988,
H. B. Glenn pers. comm.).
Encarsia sp. (as Aspidiotiphagus sp.), from Japan via Texas, imported in 1988 by H.
B. Glenn, reared on Aspidiotus nerii, 800 released in 1988 and 8,200 in 1989
against Unaspis citri, but not currently distinguishable from existing species, so
establishment uncertain (H. B. Glenn pers. comm.).
Encarsia spp., from Hong Kong and China, imported in 1986 and 1988 by R. I. Sailer
and F. D. Bennett against Dialeurodes citrifolii; was not bred in laboratory and
was not released (AD 1986, 1988).
Eretmocerus mundus Mercet, from Israel, imported by F. D. Bennett and R. Nguyen
in 1990, and from Sudan in 1991, against Bemisia tabaci (AD 1990, 1991).
Eretmocerus sp., from an unspecified country, released in 1952 from Winter Haven
(Polk County) against Dialeurodes citri, not established (Denmark 1964, Brown-
ing 1990). ~
Eretmocerus spp., from California, imported by F. D. Bennett in 1989 against Bemisia
tabaci, and 100 adults were shipped from quarantine to Bradenton (Manatee
County) and 65 to Apopka (Orange County) with release intended; from Brazil,
Costa Rica, Honduras, Mexico, and Puerto Rico, imported by F. D. Bennett in
1990, and from Brazil, Guatemala, and Venezuela, imported by F. D. Bennett in
1991 (AD 1990, 1991).









Florida Entomologist 76(1)


TABLE 2. (Continued)

Pteroptrix chinensis (Howard), from Hong Kong, imported in 1989 by H. W. Browning
for biological studies against Chrysomphalus aonidum and Aonidiella aurantii,
and no releases were made (AD 1989, H. W. Browning pers. comm.).
HYMENOPTERA: APHIDIIDAE
Aphidius gifuensis Ashmead, from Taiwan via Delaware, imported in 1972 by N. R.
Spencer as a biocontrol agent for Myzus persicae, released in a greenhouse in
Alachua County but was difficult to distinguish from native species and establish-
ment is uncertain (Denmark & Porter 1973, BIRL 1992, N. R. Spencer pers.
comm.).
Aphidius matricariae Haliday, from France, 1400 adults were released in 1957 at two
sites, and 300 adults from India were released in 1958 at two sites, against Myzus
persicae (from Quincy, Gadsden County) and Sipha flava (from Belle Glade,
Palm Beach County); not established (Denmark 1964, Charpentier et al. 1972,
Denmark & Porter 1973, Pefia 1990). ~
Aphidius picipes (Nees), from France via Delaware, imported in 1972 and 1973 by N.
R. Spencer as a biocontrol agent for Myzus persicae (BIRL 1992).
Aphidius smith Sharma & Rao, from India via New Jersey, released in 1958 at two
sites (including from Belle Glade, Palm Beach County) against Acyrthosiphon
pisum and sugarcane aphids, current status in Florida not reported (Denmark
1964, Angalet & Coles 1966, Charpentier et al. 1972, Clausen 1978, Grant &
Lambdin 1990, Jansson & Pefia 1990). ~
Aphidiussp., origin not stated, released in 1958 against Myzus persicae, not established
(Jackson 1990). ~
Binodoxys indicus Subba Rao & Sharma, from India via Delaware, imported in 1971
by A. G. Selhime as a potential biocontrol agent for Aphis gossypii (Denmark &
Porter 1973, BIRL 1992).
Praon exsoletum (Nees), from the Mediterranean, 900 adults were released in 1957 at
three sites from Belle Glade, Palm Beach County, against Siphaflava, but there
were no recoveries (Denmark 1964, Charpentier et al. 1972). ~
Trioxys complanatus Quilis, from France, 635 adults were released in 1957 at one site
(from Belle Glade, Palm Beach County) against Sipha flava, not established
(Denmark 1964, Charpentier et al 1972). ~
HYMENOPTERA: BETHYLIDAE
Goniozus indicus Ashmead, from India via Delaware, imported in 1961 by J. R. Gifford
as a potential biocontrol agent of Diatraea saccharalis; 87 cocoons were received;
although it parasitized this host in the laboratory, there were no releases (Gifford
1964, 1965, Charpentier et al. 1972, BIRL 1992).
HYMENOPTERA: BRACONIDAE
Alabagrus stigma (Brull6), from Peru, released in 1932 against Diatraea saccharalis,
established, causes 5-20% parasitism; from Argentina, imported in 1979 against
Spodoptera [even though it already was established in Florida!] (Jaynes 1938,
1939, Wilson 1941, Annand 1944, AD 1919, Bennett 1971, Charpentier et al.
1967, 1972, Hall 1986, Bennett et al. 1990, D. G. Hall pers. comm.). *
Apanteles angaleti Muesebeck, from India, released in Gadsden County in 1955-1957
against Helicoverpa and Heliothis spp., not established (Jackson 1990). ~
Apanteles diatraeae Muesebeck, from Arizona, 416 adults were released in 1934 in
Indian River County against Diatraea saccharalis, but no recoveries were made
(Scaramuzza & Ingram 1942, Gifford 1964, Charpentier et al. 1972, Bennett et
al. 1990). ~


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy 19

Apanteles sp., from Uruguay, imported in 1945-1946 against Spodoptera frugiperda
(Annand 1947).
Apanteles sp., from India via Delaware, imported in 1964 by J. R. Gifford as a biocontrol
agent for Diatraea saccharalis (BIRL 1992).
Apanteles sp., from Colombia, emerged in quarantine from Urbanus proteus L. which
was imported by R. I. Sailer in 1982 as host for Ardalus sp. (AD 1982).
Ascogaster quadridentata Wesmael, from Europe, released against Diatraea sac-
charalis (though a parasitoid of codling moth), not established (Gifford 1964). -
Biosteres arisanus (Sonan), permit issued for its importation in 1967 (as Biosteres
oophilus); from Hawaii, imported in 1974 (as Opius oophilus) by P. D. Greany
against Anastrepha suspense, 400 released in 1974 and 8,804 in 1975, in Dade
County, not established (Denmark & Porter 1973, AD 1974, Grissell 1978, Swan-
son 1978, Baranowski 1986, H. B. Glenn pers. comm.).-
Biosteres vandenboschi (Fullaway), permit issued for its importation in 1967; from India
via Hawaii, imported in 1969 (as Opius persulcatus) by R. M. Baranowski,
against Anastrepha suspense, but no culture was established and no releases
were made; from Hawaii (as Opius vandenboschi), 455 released in 1985, 777 in
1986, 720 in 1987, and 306 in 1988, in Dade County against Anastrepha suspense,
not established (Denmark & Porter 1973, Baranowski 1986, H. B. Glenn pers.
comm.). ~
Bracon kirkpatricki (Wilkinson), probably from Kenya, released in 1975 against Pec-
tinophora gossypiella, probably not established [this release may have resulted
from work in Arizona reported by Bryan et al. 1973] (Mead 1976). ~
Bracon vestiticida (Viereck), from Peru, imported in 1941 and released in Alachua
County against Anthonomus grandis, not established (Annand 1944, Berry 1947,
Clausen 1978, Gate et al. 1990). ~
Bracon sp., from India via Delaware, imported in 1964 by J. R. Gifford as a potential
biocontrol agent for Diatraea saccharalis, 50 adults were imported, but would
not parasitize this host in the laboratory (Gifford 1964, Charpentier et al. 1972,
BIRL 1992).
Uga menoni Kerrich, from Korea via Delaware, imported in 1990, released in 1990 by
L. Nong and F. D. Bennett in Alachua County as a potential biocontrol agent
for Epilachna varivestis, not established (BIRL 1992, Nong & Bennett in press).

genus and species indet., from Argentina, imported in 1988 by R. M. Baranowski as a
parasitoid of Phoridae which were contaminants of a shipment of natural enemies
of Anastrepha suspense, not released from quarantine (AD 1988, H. B. Glenn
pers. comm.).
Campyloneurus mutator F., from India via Delaware, imported in 1964 by J. R. Gifford
as a potential biocontrol agent for Diatraea saccharalis; 266 adults were im-
ported; though it was reared on D. saccharalis in the laboratory, mass-rearing
was unsuccessful and none was released (Charpentier et al. 1972, BIRL 1992).
Cardiochiles diaphaniae Marsh, from Colombia, imported in 1984 by J. E. Pefia and V.
H. Waddill, 6 females and 1 male released in Dade County in 1984 against
Diaphania hyalinata and D. nitidalis, but not established (AD 1984, Stange
1986, Jansson & Pefia 1990, J. E. Pefia pers. comm.); imported in 1989 and 1991,
124 females and 168 males released in 1992 in Dade County, recovered before
but not after hurricane Andrew (September 1992) and released in 1992 in Alachua
County by J. L. Capinera (AD 1989, 1991, Bennett & Capinera in press, J. L.
Capinera pers. comm., J. E. Pefia pers. comm.). ~
Chelonus annulipes Wesmael, from Europe; 6,000 adults were received in 1938 from
the European Corn Borer Laboratory (Moorestown, NJ) and were released in









Florida Entomologist 76(1)


TABLE 2. (Continued)

two locations against Diatraea saccharalis; not established (Gifford 1964, Char-
pentier et al. 1972). ~
Chelonus blackburni Cameron, probably from Hawaii, released in 1975 against Pec-
tinophora gossypiella, probably not established [this release may have resulted
from work in Arizona reported by Bryan et al. 1973] (Denmark & Porter 1973,
Mead 1976). ~
Chelonus busckiellaViereck, from Colombia, imported in 1991 by J. L. Capinera against
Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera pers. comm.).
Chelonus heliopae Gupta, from India, released in Gadsden County in 1955-1957 against
Heliothis virescens, not established (Denmark 1964, Jackson 1990).
Chelonus narayani Subba Rao, from India, released in Gadsden County in 1954-1957
against Helicoverpa and Heliothis spp., not established (Jackson 1990). ~
Chelonus texanus Cresson, from Bolivia, imported in 1980 against Spodoptera spp. (AD
1980).
Chelonus sp., from Colombia, imported in 1984 by J. E. Pefia, but not released (AD
1984, J. E. Pefa pers. comm.).
Cotesia flavipes Cameron, from India via Delaware, imported in 1962, released in 1963
in Palm Beach County against Diatraea saccharalis, established, achieves on
average 40-55% parasitism of the target late each season (Denmark 1964, Gifford
1964, Gifford & Mann 1967, Charpentier et al. 1972, Clausen 1978, Hall 1986,
Bennett 1970, Bennett et al. 1990, BIRL 1992). *
Cotesia plutellae Kurdjumov, from Malaysia via England, imported in 1990 by R. K.
Jansson and F. D. Bennett as a potential biocontrol agent for Plutella xylostella,
<200 released in 1990 in Dade County by R. K. Jansson, and 118 in Seminole
County by G. L. Leibee, not established; from southeast Asia via a commercial
producer of biocontrol agents in Texas, large numbers released by a vegetable
grower in Orange County in 1990; from southeast Asia via Hawaii, Washington
and then via a commercial producer of biocontrol agents in Texas, >20,000 re-
leased in experimental plots of vegetables in Lake County in 1992 by E. R.
Mitchell & J. R. McLaughlin in 1992, too recently to determine whether establish-
ment occurred (Jansson & Pefia 1990, G. L. Leibee pers. comm., E. R. Mitchell
& J. R. McLaughlin pers. comm.). ?
Diachasmimorpha longicaudata (Ashmead), from Mexico and Hawaii, 200 released in
1972 (in Dade County), 146,637 in 1973 (in Broward, Collier, Dade, Highlands,
Indian River, Lee, Manatee, Martin, Monroe and Palm Beach counties),
1,006,413 in 1974 (in Brevard, Broward, Charlotte, Collier, Dade, Glades, Hen-
dry, Hillsborough, Lee, Manatee, Martin, Monroe, Okeechobee, Palm Beach,
Polk, Sarasota, St. Lucie and and Volusia counties), 18,200 in 1975 (in Dade,
Collier and Orange counties), 1,350 in 1976 (in Dade County), and 1,600 in 1977
(in Dade County) against Anastrepha suspense, established, causing reduction
of target populations by 40% (Denmark & Porter 1973, Grissell 1978, Baranowski
1979, 1986, Swanson 1982, Thompson 1989, Browning 1990, H. B. Glenn pers.
comm.). *
Diachasmimorpha tryoni (Cameron), from Hawaii (as Biosteres tryoni), 579 released
in 1982, 239 in 1983, 400 in 1984, 7,709 in 1985, 16,880 in 1986 and 4,815 in 1987,
by H. B. Glenn, all in Dade County against Anastrepha suspense, recovered,
perhaps established (Denmark & Porter 1973, ROBO 1982, Baranowski 1986, H.
B. Glenn pers. comm.). *
Digonogastra amabilis (Br6thes), perhaps from Argentina, released against Diatraea
saccharalis, date of importation and county of release not specified, not estab-
lished (Gifford 1964). ~


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy 21

Digonogastra kimballi Kirkland, from Mexico via Missouri, about 200 adults released
in 1981, released in 1981 against Diatraea saccharalis in Palm Beach County by
O. Sosa, but no recoveries were made and probably is not established (ROBO
1981, Bennett et al. 1990 [as Iphiaulax sp.], O. Sosa pers. comm., D. G. Hall
pers. comm.). ~
Digonogastra rimac (Wolcott), from Peru, in 1932 12,984 adults were released at two
sites, and in 1936 over 2,000 in 1936 were released at two sites (189 in Indian
River County and 2,032 in Palm Beach County), against Diatraea saccharalis; a
very few individuals were recovered in the field in 1934 (2 years after the first
release), but permanent establishment has not occurred (Jaynes 1938, 1938, Gif-
ford 1964, Charpentier et al. 1967, 1972, Oatman & Clausen 1978, Bennett 1971,
Bennett et al. 1990). ~
Doryctobracon areolatus (Szepligeti), from Trinidad (as by R. M. Baranowski and F.
D. Bennett against Anastrepha suspense, but material received from Brazil in
1989 belonged to other species such as D. areolatus, etc. (F. D. Bennett pers.
comm., H. B. Glenn pers. comm.).
Doryctobracon crawfordi (Viereck), permit issued for its importation and release in
1968 (as Parachasma crawfordi); from Ecuador, 11 adults were collected in
Ecuador in 1986 by F. D. Bennett, with the possibility of their use against
Anastrepha suspense, but was not cultured successfully and no release was made
(Denmark & Porter 1973, H. B. Glenn pers. comm.).
Doryctobracon fluminensis (Costa Lima), from Colombia, imported in 1984 by R. M.
Baranowski and J. E. Pefia against Anastrepha suspense, but not cultured in
the laboratory and not released (AD 1984, Stange 1986, H. B. Glenn pers.
comm.).
Doryctobracon trinidadensis Gahan, permit issued for its importation in 1968 (as
Parachasma trinidadense); from Trinidad, 23 released in 1975, 86 in 1978, 731
in 1985, 2,036 in 1986, and 803 in 1987, in Dade County (as Opius trinidadense)
against Anastrepha suspense, not recovered since 1986, presumably not estab-
lished (Denmark & Porter 1973, Swanson 1978, Baranowski 1986, H. B. Glenn
pers. comm.). ~
Glyptapanteles flavicoxis (Marsh), from India via Delaware, imported at an unspecified
date; its original host was Lymantria obfuscata Walker, and it was imported
against Lymantria dispar (BIRL 1992).
Glyptapanteles sp. probably caffreyi (Muesebeck), from Ecuador, imported in 1982-1983
by V. H. Waddill, 2,025 released in 1982-1983 in Dade County against Anticarsia
gemmatalis, current status unknown (Boethel & Orr 1990). ~
Habrobracon brevicornis Wesmael, from India (as Bracon brevicornis), released in
Gadsden County in 1955-1957 against Helicoverpa and Heliothis spp., also re-
leased elsewhere against Diatraea saccharalis, not established (Gifford 1964,
Jackson 1990). ~
Hypomicrogaster diaphaniae Muesebeck, from Colombia, imported in 1984 by J. E.
Pefia, as a potential biocontrol agent for Diaphania hyalinata and D. nitidalis,
but never released (Jansson & Pefa 1990, J. E. Pefa pers. comm.).
Hypomicrogaster sp., from Colombia, imported in 1991 by J. L. Capinera against
Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera pers. comm.).
Iphiaulax sp. [perhaps Rhaconotus roslinensis Lal), from India, 134 adults were im-
ported in 1965, but they failed to parasitize the target (Diatraea saccharalis) in
the laboratory, and no releases were made (Charpentier et al. 1972, F. D. Ben-
nett pers. comm.).
Lipolexis scutellaris Mackauer, from Malaysia, imported in 1991 by S. Watts against
Toxoptera aurantii [and as a precaution lest T. citricida (Kirkaldy) should arrive
in Florida], and sent to Dade County (AD 1991).









Florida Entomologist 76(1)


TABLE 2. (Continued)

Meteorus laphygmae Viereck, from Colombia via Missouri, imported in 1975 by T. R.
Ashley against Spodoptera spp., but died in quarantine (AD 1975).
Meteorus sp., from Paraguay and Peru, imported in 1979 against noctuids (AD 1979).
Microplitis demolitor (Wilkinson), from Australia, released in 1985 against Helicoverpa
and Heliothis spp. and Pseudoplusia includes (Powell 1989, Boethel & Orr
1990). ~
Microplitis manilae (Ashmead), from Thailand, imported by M. Shepard in 1981, 5,000
released in 1982 in Dade County against Spodopterafrugiperda by V. H. Waddill,
not established (ROBO 1982, Gross & Pair 1986, V. H. Waddill pers. comm.). -
Microplitis rufiventris Kokujev, from Egypt, released in 1983-1984 in Dade County by
S. D. Pair and V. H. Waddill against Spodoptera frugiperda, not established
(Gross & Pair 1986). ~
Myosoma chinensis Sz6pligeti, from India via Delaware, imported in 1963 by J. R.
Gifford (as Bracon chinensis) and 1964 by G. A. Mann (a total of 176 adults) as
a biocontrol agent for Diatraea saccharalis, but would not parasitize this host in
the laboratory and was not released (Gifford 1964, Charpentier et al. 1972, BIRL
1992).
Opius bellus Gahan, from Trinidad, imported in mid-1970s from Trinidad as a potential
biocontrol agent of Anastrepha suspense, not released; from Brazil, imported by
R. M. Baranowski and F. D. Bennett in 1989 with other material, but only 1
female was in the shipment and was not released (Swanson 1978, Baranowski
1986, AD 1989, F. D. Bennett pers. comm., H. B. Glenn pers. comm.).
Phaenocarpa anastrephaeMuesebeck, from Brazil, 1 female was imported accidentally
in 1989 by R. M. Baranowski and F. D. Bennett in a shipment of other material,
but was not released (F. D. Bennett pers. comm., H. B. Glenn pers. comm.).
Protomicroplitis sp., from Paraguay and Peru, imported in 1979 against noctuids (AD
1979).
Psyttalia concolor Sz6pligeti, from R6union via France and Delaware, imported by R.
M. Baranowski, 127 released in 1978 (Dade County), 8,618 released in 1979 (Dade
County), 22,410 in 1980 (19,335 in Dade County, 135 in Broward County, 318 in
Citrus County, 481 in Hillsborough County, 1,435 in Palm Beach County, 318 in
Pasco County, and 388 in Polk County), and 1,783 in 1983, against Anastrepha
suspense, established (Swanson 1982, Baranowski 1986, BIRL 1992, H. B. Glenn
pers. comm.). *
Psyttalia fletcheri (Silvestri), permit issued in 1968 for its importation; from Hawaii,
1,755 released in 1988 in Dade County against Anastrepha suspense, not estab-
lished (Denmark & Porter 1973, H. B. Glenn pers. comm.). ~
Psyttalia incisi (Silvestri), from Hawaii, 236 released in 1983, 438 in 1985, and 2,160 in
1988, all in Dade County against Anastrepha suspense, probably not established
(Baranowski 1986, H. B. Glenn pers. comm.). ~
Rhaconotus signipennis Walker, from India via Delaware, 129 adults imported in 1964
by G. A. Mann and J. R. Gifford as a potential biocontrol agent for Diatraea
saccharalis, but no releases were made (Gifford 1964, Charpentier et al. 1972,
BIRL 1992).
Rhaconotus sp., from India via Delaware, imported in 1964 against Diatraea sac-
charalis by G. A. Mann (BIRL 1992).
Stantonia lamprosemae Muesebeck, from Colombia, imported in 1991 by J. L. Capinera
against Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera pers.
comm.).
Stenobracon deesae (Cameron), from India via Delaware, 497 adults were imported in


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy


1964 against Diatraea saccharalis by J. R. Gifford, but no releases were made
(Gifford 1964, BIRL 1992).
Stenobracon nicevillei Bingham, from India via Delaware, 171 adults were imported in
1964 against Diatraea saccharalis, by J. R. Gifford and G. A. Mann; this host
was parasitized in the laboratory, but no releases were made (Gifford 1964, Char-
pentier et al. 1972, BIRL 1992).
Triaspis vestiticida Viereck, from Peru, imported in 1941 against Anthonomus grandis,
released in Alachua County but not established (Annand 1944, Berry 1947,
Clausen 1978, Cate et al. 1990).
Urosigalphus schwarzi Gibson, from Guatemala, imported in 1974 by W. H. Whitcomb
against Anthonomus grandis, but died in quarantine (AD 1974).
Utetes anastrephae (Viereck) (as Opius anastrephae and Bracanastrepha anastrephae),
permit issued for its importation in 1968; from Brazil, a shipment of other mater-
ial imported by R. M. Baranowski and F. D. Bennett in 1989 happened to have
3 females and 1 male, which were not released, in part because the species was
found in 1973 apparently as an immigrant to Florida and parasitoid of Anastrepha
suspense (Denmark & Porter 1973, Frank & McCoy 1992, F. D. Bennett pers.
comm., H. B. Glenn pers. comm.).
Zelomorpha sp., from Costa Rica, imported in 1982 by M. Shepard, G. Carner, and V.
H. Waddill, 6 released in Dade County in 1982 (recoveries were made early in
1983) and in 1983 and 1985 against Anticarsia gemmatalis, current status un-
known (ROBO 1982, Boethel & Orr 1990, V. H. Waddill pers. comm.). ~
genus and species indet. (subfamily Opiinae), from Argentina, imported in 1988 by R.
M. Baranowski against Anastrepha suspense, but not released from quarantine
(AD 1988, H. B. Glenn pers. comm.).
HYMENOPTERA: CHALCIDIDAE
Brachymeria sp., from Colombia, imported in 1984 by J. E. Pefia as a potential biocon-
trol agent for Diaphania hyalinata and D. nitidalis, but never released (Jansson
& Pefia 1990, J. E. Pefia pers. comm.).
Dirhinus giffardii Silvestri, from R4union via Delaware, imported in 1978, against
Anastrepha suspense, 1,750 released in 1978 in Dade County, 17,040 released in
1979 in Dade County, 66,748 released in 1980 (59,185 in Dade County, 1,330 in
Broward County, 105 in Collier County, 171 in Hernando County, 1,484 in Hill-
sborough County, 35 in Lee County, 35 in Martin County, 2,200 in Monroe
County, 1,690 in Palm Beach County, 171 in Pasco County, and 342 in Pinellas
County), and 4,000 in 1986 by R. M. Baranowski, not established (Swanson 1982,
Baranowski 1986, BIRL 1992, H. B. Glenn pers. comm.).
Dirhinus himalayanus Westwood, from Morocco via France, imported in 1990 against
Musca domestic by P. B. Morgan for laboratory evaluation (AD 1990, P. B.
Morgan pers. comm.).
Spilochalcis diaphaniae (Ashmead), from Colombia, imported in 1984 by J. E. Pefia
against Diaphania hyalinata and D. nitidalis, but never released (Jansson &
Pefia 1990, J. E. Pefia pers. comm.).
Spilochalcis sp., from Colombia, imported in 1991 by J. L. Capinera and H. A. Smith
against Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera pers.
comm.).
HYMENOPTERA: DIAPRIIDAE
Basalys sp., from Mauritius, an unsolicited shipment in 1974 received by R. S. Patter-
son against Stomoxys calcitrans (AD 1974).
Coptera merceti (Say), France, imported in 1986, 1987, and 1988 by P. B. Morgan and
R. S. Patterson against Musca domestic and Stomoxys calcitrans; from Hun-









Florida Entomologist 76(1)


TABLE 2. (Continued)

gary, imported in 1986 and 1988; all for laboratory study, without release (AD
1986, 1987, 1988, P. B. Morgan pers. comm.).
Trichopria cilipes Kieffer, from France, imported in 1990 against Stomoxys calcitrans,
all for laboratory study, without release (AD 1990, P. B. Morgan pers. comm.).
Trichopria painter Huggert & Morgan, from Zimbabwe, imported in 1986 together
with T. cilipes, described later, used for laboratory study, without release (P.
B. Morgan pers. comm).
Trichopria stomoxydis Huggert, from Mauritius, imported in 1981 and 1983 by P. B.
Morgan against Stomoxys calcitrans; from Hungary and from Zimbabwe, im-
ported in 1986 by R. S. Patterson against Musca domestic and Stomoxys calci-
trans; from Mauritius via France, imported in 1988-1989 by P. B. Morgan and D.
R. Barnard; all for laboratory study, without release (AD 1981, 1983, 1986, 1988,
1989, Stange 1986, P. B. Morgan pers. comm.).
HYMENOPTERA: DRYINIDAE
Pseudogonatopus variistriatus Fenton, from Jamaica via Trinidad and New Jersey,
imported in 1959 and released from Belle Glade (Palm Beach County) against
Saccharosydne saccharivora, not established (Bennett 1960, Denmark 1964, F.
D. Bennett pers. comm.) [note that a native species, P. arizonicus Perkins, has
been reported from S. saccharivora in Florida (Gifford 1964, Olmi 1984)]. ~
HYMENOPTERA: ENCYRTIDAE
Anagyrus antoninae Timberlake, from Japan via Texas, released in 1950 in Palm Beach
SCounty (from Belle Glade), in 1954 in Hendry County, in 1955 in Palm Beach and
Martin counties, and in 1957 in Palm Beach County against Antonina graminis,
established, but not found in recent surveys and possibly displaced by Neodus-
metia sangwani and Pseudectroma sp. (Questel & Genung 1957, 1961, Denmark
1964, Bartlett 1978, Frank 1990, Bennett & Capinera in press, Bennett in this
symposium). *
Anagyrus diversicornis Mercet, from Europe, released from Belle Glade (Palm Beach
County), by USDA-ARS, date unspecified, against Antonina graminis, not es-
tablished (Denmark 1964).
Comperiella bifasciata Howard, from Asia via California, imported in 1926 by J. R.
Watson against Chrysomphalus aonidum and released in Polk County (Watson
1926).
Copidosoma floridanum (Ashmead), from Argentina via Delaware, against Anticarsia
gemmatalis and Pseudoplusia includes (AD 1975).
Copidosoma sp., origin not stated, released from Quincy (Gadsden County) against
Trichoplusia ni, year of release not stated (Denmark 1964). ~
Hambletonia pseudococcina Compere, from South America via Puerto Rico, imported
in 1943 but received dead, imported and released in Highlands, Palm Beach, and
St. Lucie counties in 1944 against Dysmicoccus brevipes, became established
first in Highlands County (Clausen 1956, Bartlett 1978). *
Leptomastidea abnormis (Girault), from Italy via California, released under the name
Leptomastix abnormis in Pinellas County in 1917 against Planococcus citri, and
recorded in 1923 from that county (Watson 1924); the current widespread distri-
bution of the species in eastern North America may have resulted from its much
earlier immigration with host material (Bartlett 1978). *
Leptomastix dactylopii Howard, from the neotropics via California, imported in 1939
and released in 1940 in Sarasota County against Planococcus citri (Watson &
Thompson 1940, 1941); the species is believed to be native to Brazil though dis-


March, 1993










Insect Behavioral Ecology-'92: Frank & McCoy


tribute widely in the West Indies and parts of the southern USA [and may thus
have occurred in Florida long before this release] (Bartlett 1978). *
Metaphycus helvolus (Compere), from Africa via California, fewer than 2,000
parasitoids were released in 1970-1971 against Saissetia oleae (Bernard), not
established, but then it was discovered that the intended target in Florida was
S. neglect (rather than S. oleae, even though S. oleae is widespread) (Denmark
& Porter 1973, McCoy 1985, Browning 1990, F. D. Bennett pers. comm.). ~
Metaphycus luteolus Timberlake, permit issued in 1971 for its release in Florida, pre-
sumably against Coccus hesperidum (Denmark & Porter 1973).
Neodusmetia sangwani (Subba Rao), from India via New Jersey, released in 1957 (from
Belle Glade) in Palm Beach County against Antonina graminis, with further
releases in the same county in 1959 via Texas, established and widespread (De-
nmark 1964, Bennett & Capinera in press). *
Ooencyrtus kuvanai (Howard), from Japan via northeastern USA, released in 1971 in
Santa Rosa County against Lymantria dispar (Denmark & Porter 1973, Poucher
1974) [spellings kuwanai, kuwanae, kuwani, and kuvanae in assorted publica-
tions are erroneous]. ~
Ooencyrtus submetallicus (Howard), from Montserrat via Trinidad and Delaware, im-
ported in 1973 and 1974 by N. R. Spencer and W. H. Whitcomb as a potential
biocontrol agent for Nezara viridula, but occurs already in Florida (AD 1973,
1974, BIRL 1992, F. D. Bennett pers. comm., N. R. Spencer pers. comm.).
Pseudaphycus mundus Gahan, from Louisiana, released in 1932, 1934, and 1936 in Palm
Beach, Hendry, and Indian River counties against Dysmicoccus boninsis, D.
brevipes, and Saccharicoccus sacchari, under the mistaken belief that it was
Aphycus terryi Fullaway (imported to Louisiana from Hawaii), established
(Bynum 1937, Gahan 1946, Charpentier et al. 1972, Bartlett 1978, Bennett et al.
1990). *
Pseudectroma europaea (Mercet), from France, released in 1957 and 1959 (from Belle
Glade) in Palm Beach County against Antonina graminis, not established (Den-
mark 1964, Bennett & Capinera in press). ~
Tachinaephagus stomoxicida Subba Rao, from Mauritius via California, imported in
1981, 1982 and 1983 by P. B. Morgan and R. S. Patterson against Stomoxys
calcitrans; from Mauritius via France, imported in 1989-1990 by D. R. Barnard
and P. B. Morgan; the imported material was used only for laboratory study,
without release (AD 1981, 1982, 1983, 1989, 1990, P. B. Morgan pers. comm.).
Tachinaephagus zealandicus Ashmead, permit issued in 1971 for its importation and
release in Florida; from Mauritius, imported in 1975 by P. B. Morgan against
Stomoxys calcitrans; from Mauritius via France, imported in 1989 by D. R.
Barnard; this species is adventive in the USA and was reported from Florida by
Butler et al. (1981); the imported material was used only for laboratory study,
without release (AD 1975, 1989, Denmark & Porter 1973, P. B. Morgan pers.
comm.).
Thomsonisca sankarani Subba Rao, from India, imported in 1978 by G. R. Buckingham
(from Pseudaulacaspis barber Green), 40 released in Dade County against
Pseudaulacaspis cockerelli by R. M. Baranowski (AD 1978, F. D. Bennett pers.
comm., G. R. Buckingham pers. comm., H. B. Glenn pers. comm.).
genus and species indet., from Brazil, imported in 1989 by P. B. Morgan against
Stomoxys calcitrans, used for laboratory study and not released (AD 1989, P.
B. Morgan pers. comm.).
HYMENOPTERA: EUCOILIDAE
Trybliographa daci Weld, from the Indo-Australian region via France, imported by R.
M. Baranowski in 1977 against Anastrepha suspense, 6,259 released in 1979 (all









26 Florida Entomologist 76(1) March, 1993

TABLE 2. (Continued)

in Dade County), 37,016 released in 1980 (925 in Broward County, 105 in Collier
County, 34,634 in Dade County, 800 in Hillsborough County, and 552 in Monroe
County) by R. W. Swanson and others, established (ROBO 1981, Baranowski
1986, Glenn & Baranowski 1987, BIRL 1992, H. B. Glenn pers. comm.). *
Trybliographa sp. (as Ganaspis sp.), from Brazil, imported in 1989 by F. D. Bennett
against Anastrepha suspense (AD 1989).
HYMENOPTERA: EULOPHIDAE
Aceratoneuromyia indica (Silvestri), permit for importation issued in 1967; from Costa
Rica, imported in mid-1970s by R. M. Baranowski and R. W. Swanson against
Anastrepha suspense; from Colombia, imported in 1984 by R. M. Baranowski
and J. E. Pefia, 4,000 released in Dade County, recovered on 4 separate occasions
and believed to be established; received from Texas in 1986, 2,000 released in
1986, 2,100 in 1987, all in Dade County (Denmark & Porter 1973, AD 1984, H.
B. Glenn pers. comm.). *
Ardalus sp., from Colombia, imported in 1982 by R. I. Sailer, original host Urbanus
proteus L., so presumably against some pest hesperiid, died in quarantine (AD
1982).
Dahlbominus fuscipennis (Zetterstedt), permit issued in 1971 for its importation and
release in Florida; from Europe via the northeastern USA, released by R. C.
Wilkinson in 1971 against Neodiprion lecontei, not established (Denmark & Por-
ter 1973, A. T. Drooz pers. comm., R. C. Wilkinson pers. comm.). ~
Edovum puttleri Grissell, from Colombia via Missouri, imported in 1983 by R. I. Sailer
against Leptinotarsa decemlineata, 30 released in Alachua County in 1983 by B.
Munir, current status unknown (ROBO 1983, Stange 1986, Jansson & Pefa 1990). ~
Euplectrus sp. nr. comstocki Howard (as plathypenae Ashmead), from Colombia via Mis-
souri, imported in 1975 by T. R. Ashley against Spodoptera sp., released in 1976 as
a biocontrol agent for Anticarsia gemmatalis (AD 1975, Grissell 1978). ~
Euplectrus puttleri Gordh, from Brazil, imported by B. Puttler and V. H. Waddill and
released in 1981-1982 in Collier (90), Dade (133), and Palm Beach counties (242),
against Anticarsia gemmatalis, established (ROBO 1981, 1982, Boethel & Orr
1990, V. H. Waddill pers. comm.). *
Goetheana shakespearei Girault, from Africa via Puerto Rico, imported by J. R. Watson
(under the name Dasyscapus parvipennis Gahan) in 1939 against Selenothrips
rubrocinctus, 500 puparia without record of release; from Africa via California,
imported under the name Goetheana parvipennis (Gahan), about 40 adults were
released in 1986 in Dade County in a greenhouse against S. rubrocinctus; from
Africa via Puerto Rico, about 40 adults were released against S. rubrocinctus in
Dade County; pupae were found in 1992 in Dade County at a location far from
the release sites, so evidently established, but not clearly as a result of the
releases, perhaps independently as an immigrant (Watson & Thompson 1940,
Mizell & Tedders 1990, Bennett et al. 1993, H. Glenn & F. D. Bennett pers.
comm.). *
Horismenus elineatus Schauff, from Bolivia via Texas, imported against Elasmopalpus
lignosellus, about 450 released in Gadsden County and about 200 in Jackson
County by J. E. Funderburk in 1990, not established; from Bolivia via Texas and
Hawaii, imported by D. G. Hall and F. D. Bennett in 1991, about 7,500 released
in 1992 in Palm Beach and Hendry counties, too recently to determine whether
establishment occurred (D. G. Hall pers. comm., J. E. Funderburk pers. comm.). ?
Horismenus sp., from Mexico via Texas, imported in 1976 against Anthonomus grandis
(AD 1976).










Insect Behavioral Ecology-'92: Frank & McCoy 27

Pediobius facialis (Giraud), from Japan via Delaware, imported in 1982 by R. I. Sailer,
released in 1981-1982 in Dade County (1,000) by V. H. Waddill against Tricho-
plusia ni (ROBO 1981, 1982 BIRL 1992).
Pediobius foveolatus (Crawford), permit issued for its importation and release in 1968;
from India via Maryland, imported in 1973 and 1974 via Maryland by N. R.
Spencer against Epilachna varivestis, about 1,500 released in 1974 in Gadsden
County and 1,000 in Alachua County, released in 1975-1976 resulting in 3 years
of reduced populations of the target, though the parasitoid does not overwinter
in northern Florida so needs reintroduction each year; from Japan via Delaware,
imported in 1982 by R. I. Sailer for study (AD 1973, 1974, 1982, Denmark &
Porter 1973, Grissell 1978, Sailer 1981, Boethel & Orr 1990, Jansson & Pefia
1990, BIRL 1992, N. R. Spencer pers. comm.). ~
Tetrastichus ceroplastae Girault, permit issued in 1969 for its importation into Florida
[this species is recorded from the Indian Ocean islands of Madagascar and
Mauritius as a parasitoid of scale insects] (Denmark & Porter 1973).
Tetrastichus fennahi Schauff, from Puerto Rico, imported in 1977 but died during ship-
ping or in quarantine; from the Dominican Republic, imported in 1990 by H. W.
Browning against Diaprepes abbreviatus, but no culture was established and no
releases were made (AD 1977, 1990, H. W. Browning pers. comm.).
Tetrastichus gala Walker, from the Dominican Republic, imported in 1990 by H. W.
Browning against Diaprepes abbreviatus, but no culture was established and no
releases were made (AD 1990, H. W. Browning pers. comm.).
Tetrastichus giffardianus Silvestri, permit issued in 1968 for its importation into Florida
against Anastrepha suspense [occurs in Hawaii] (Denmark & Porter 1973).
Tetrastichus haitiensis Gahan, from Puerto Rico, released in 1969 in Palm Beach
County, in 1970 in Orange County, and in 1971 in Palm Beach County, against
Diaprepes abbreviatus, at first did not establish, but established by 1978 (Sutton
et al. 1972, Denmark & Porter 1973, Beavers & Selhime 1975, Beavers et al.
1980, Woodruff 1981), and recovered from eggs of the host in Miami in 1986
(Bennett et al. 1990); from Puerto Rico, imported in 1977 but died during shipping
or in quarantine, and in 1984 and in 1990 against Diaprepes abbreviatus (AD
1977, 1984, 1990); from the Dominican Republic, imported in 1990 (mixed with
T. fennahi and hyperparasitoids) by H. W. Browning against Diaprepes ab-
breviatus, and 16 adults were shipped from quarantine to Lake Alfred (Polk
County), but it was not possible to maintain a culture on laboratory-cultured D.
abbreviatus, and no releases were made; unclear whether its presence in Miami
is the result of introduction or of immigration (AD 1990, H. W. Browning pers.
comm). *
Tetrastichus sp., from Jamaica via Trinidad, sent to New Jersey in 1959 with the
intention that it be forwarded to Florida against Saccharosydne saccharivora,
but without record of receipt in Florida and without record of release in Florida
(Bennett 1960, F. D. Bennett pers. comm.).
HYMENOPTERA: FORMICIDAE
Formica integra Nylander, from west-central Georgia, released in an ecological exper-
iment in 1973 in Alachua County, not for biocontrol purposes, destroyed by native
ants (Camponotus) (Wilkinson et al. 1980). ~
Solenopsis (Labauchena) daguerrei Santschi, from Argentina, imported in 1987 by D.
P. Wojcik against Solenopsis invicta, not released (AD 1987, D. P. Wojcik pers.
comm.).
Solenopsis (Labauchena) sp., from Brazil, imported in 1986-1987 and 1989 by D. P.
Wojcik against Solenopsis invicta, not released (AD 1986, 1987, 1989, D. P.
Wojcik pers. comm.).










28 Florida Entomologist 76(1) March, 1993

TABLE 2. (Continued)
HYMENOPTERA: ICHNEUMONIDAE
Agrypon caribbaeum Bland, from St. Croix (US-VI), imported in 1983 and brought to
Homestead (Dade County) in 1984 as a potential biocontrol agent for Diaphania
hyalinata and D. nitidalis, but not released (Jansson & Pefia 1990, J. E. Pefia
pers. comm.).
Bathyplectus anurus (Thomson), from France via Delaware, imported in 1977 by D. B.
Bouk (graduate student, Univ. Florida) as a biocontrol agent for Hyperapostica;
via Kentucky and Delaware, imported in 1978 by R. I. Sailer; via Kentucky and
Michigan, imported by R. I. Sailer in 1983 and 1,800 released in 1983 in Alachua
County by B. Munir and R. I. Sailer; probably not established (AD 1983, ROBO
1983, Munir & Sailer 1984, Stange 1986, Grant & Lambdin 1990, BIRL 1992). ~
Campoletis flavicincta (Ashmead), from Argentina, imported in 1979 against Spodopt-
era spp., Anticarsia gemmatalis, etc. (AD 1979).
Campoletis grioti (Blanchard), from Uruguay, being screened in quarantine in 1978-1980
against Spodoptera frugiperda (Stange 1982).
Casinaria sp., from Colombia, imported in 1991 by J. L. Capinera and H. A. Smith
against Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera pers.
comm.).
Aubertillus alternecoloratus (Cushman), from India via Delaware, imported in 1964 by
J. R. Gifford as a potential biocontrol agent for Diatraea saccharalis; 266 adults
were received; although it parasitized D. saccharalis in the laboratory, no individ-
uals were released (Gifford 1964, Charpentier et al. 1972, BIRL 1992).
Centeterus sp., from India via Delaware, imported in 1964 by J. R. Gifford as a potential
biocontrol agent for Diatraea saccharalis (BIRL 1992).
Coccygomimus sp., from India, in 1964-1965, 125 adults were received as potential
biocontrol agents for Diatraea saccharalis; there were no releases (Charpentier
et al. 1972).
Corsoncus magus (Cresson), from Costa Rica, 35 released in 1982 by M. Shepard, G.
Carner and V. H. Waddill in Dade County against Anticarsia gemmatalis, cur-
rent status unknown (ROBO 1982, Boethel & Orr 1990, V. H. Waddill pers.
comm.). ~
Eiphosoma dentator (F.), from Colombia, imported in 1991 by J. L. Capinera and H.
A. Smith against Diaphania hyalinata and D. nitidalis (AD 1991, J. L. Capinera
pers. comm.).
Eiphosoma vitticole Cresson, from Bolivia, imported in 1978 and 1980 by G. R. Buckin-
gham against Spodopterafrugiperda, released in 1980 in Dade County, not estab-
lished (AD 1978, 1980, Ashley et al. 1982, Stange 1982, G. R. Buckingham pers.
comm). ~
Enicospilus merdarius (Gravenhorst), from Uruguay, imported in 1945-1946 against
Spodoptera frugiperda, though this species is native to Florida (Annand 1947).
Enicospilus sp., from Paraguay and Peru, imported in 1979 against noctuids, died in
quarantine (AD 1979).
Exenterus amictorius (Panzer), from Europe via North Carolina, released in 1969
against Neodiprion lecontei, not established (Wilkinson 1969, Denmark & Porter
1973, R. C. Wilkinson pers. comm.).
Exeristes roborator (F.), a palearctic species, released against Diatraea saccharalis,
date and county not specified, not established (Gifford 1964). ~
Microcharops anticarsiae Gupta (as M. bimaculata (Ashmead)), from Colombia, Brazil,
Argentina and Costa Rica, imported in 1975 and 1980-1982 by M. Shepard, G.
Carner and V. H. Waddill, stock from Costa Rica released in 1982 by V. H.










Insect Behavioral Ecology-'92: Frank & McCoy


Waddill in Dade (5,040) and Palm Beach (60) counties, and in 1983 by L. Douthit
in Broward (485) and Dade (3,687) counties, and this or other stock in 1984-1986,
against Anticarsia gemmatalis, current status unknown (ROBO 1982, 1983,
Gupta 1987, Boethel & Orr 1990, V. H. Waddill pers. comm.). ~
Netelia sp., from Paraguay and Peru, imported in 1979 against noctuids, died in quaran-
tine (AD 1979).
Pleolophus basizonus (Gravenhorst) permit issued in 1971 for its importation and re-
lease; from Europe via North Carolina, released against Neodiprion lecontei in
1971, not established (Denmark & Porter 1973, A. T. Drooz pers. comm., R. C.
Wilkinson pers. comm.). ~
Polycyrtus sp., from Colombia, imported in 1984 by J. E. Pefa and V. H. Waddill as
a potential biocontrol agent for Diaphania hyalinata and D. nitidalis, but never
released (AD 1984, Jansson & Pefia 1990, J. E. Pefa pers. comm.).
Thymebatis sp., from Paraguay and Peru, imported in 1979 against noctuids, died in
quarantine (AD 1979).
genus and species indet.,from India via Delaware, imported in 1964 by J. R. Gifford as
a potential biocontrol agent for Diatraea saccharalis (BIRL 1992).
genus and species indet., from Brazil, imported in 1976 by N. R. Spencer against
Anticarsia gemmatalis, not released (AD 1976, N. R. Spencer pers. comm.).
HYMENOPTERA: MYRMARIDAE
Anagrus armatus Ashmead, from Jamaica via Trinidad and New Jersey, 1,014 adults
were released in 1959 at nine sites from Belle Glade (Palm Beach County) against
Saccharosydne saccharivora, but probably already occurred in Florida (Bennett
1960, Denmark 1964 [as Anagyrus sp.], Gifford 1964, Simmonds 1969, Charpen-
tier et al. 1972, Bennett et al. 1990, F. D. Bennett pers. comm.).
Erythmelus sp., from Venezuela, imported in 1991 by D. G. Hall, F. D. Bennett, and
R. Nguyen as a potential natural enemy of Leptodictya tabida, but did not repro-
duce in captivity and was not released (AD 1991, Nguyen & Hall 1991, F. D.
Bennett pers. comm., D. G. Hall pers. comm.).
HYMENOPTERA: PERGIDAE
Heteroperryia hubrichi Malaise, from Brazil, imported into quarantine in 1989-1991 as
a potential biocontrol agent for Schinus terebinthifolius by D. H. Habeck and F.
D. Bennett, but no releases were made (AD 1989, 1990, 1991, Habeck et al. in
press, D. H. Habeck pers. comm.).
Lophyrotoma zonalis (Rohwer), from Australia, imported into quarantine in 1992 as a
potential biocontrol agent for Melaleuca quinquenervia, but no releases have
been made (Habeck et al. in press, D. H. Habeck pers. comm.).
HYMENOPTERA: PERILAMPIDAE
Perilampus sp., from Colombia, imported in 1991 by J. L. Capinera with parasitoids
of Diaphania hyalinata and D. nitidalis, but is a hyperparasitoid and was not
released (AD 1991, J. L. Capinera pers. comm.).
HYMENOPTERA: PLATYGASTERIDAE
Amitus hesperidum Silvestri, from India via Mexico (in part via Texas), by A. G.
Selhime, about 22,000 parasitoids were released in 1976 in Broward County
against Aleurocanthus woglumi, established and highly successful in reducing
target populations (Grissell 1978, Hart et al. 1978, Dowell et al. 1979, Selhime
et al. 1982, McCoy 1985, Nguyen 1988, Browning 1990); with additional importa-
tion from Mexico and release in Brevard (11,286), Lee 5,102), and Martin (2,489)
counties in 1981 by W. Grandberry (ROBO 1981), and subsequent releases in
Collier, Dade, Highlands, Hillsborough, Indian River, Manatee, Monroe,









Florida Entomologist 76(1)


TABLE 2. (Continued)

Okeechobee, Palm Beach, Pinellas, Sarasota, and St- Lucie counties (Nguyen
1988). *
Amitus sp., from Puerto Rico, imported in 1990 by F. D. Bennett against Bemisia
tabaci, released in 1990 in Alachua, Collier, Dade and Manatee counties, and
specimens were captured in the field up to a year after release (F..D. Bennett
pers. comm.). ?
Amitus sp., from Guatemala, imported in 1991 by F. D. Bennett against Bemisia tabaci,
but was not released (AD 1991).
Fidiobia citri (Nixon), from Jamaica and California, imported in 1989 by H. W. Brown-
ing against Diaprepes abbreviatus and Asynonychus godmanni, but culture died
and no releases were made (AD 1989, H. W. Browning pers. comm.).
Platystasius asinus Loiacono, from Chile, imported in 1990 from a laboratory culture
against Diaprepes abbreviatus and other weevil pests of citrus bh H. W. Brown-
ing; 22 adults were shipped from quarantine to Lake Alfred (Polk County), re-
leased in 1991 but establishment yet uncertain (AD 1990, H. W. Browning pers.
comm.). ?
HYMENOPTERA: PTEROMALIDAE
Catolaccus grandis (Burks) (as Heterolaccus), from Mexico, imported in 1974 by W. H.
Whitcomb against Anthonomus grandis, for trans-shipment to Texas, but died
in quarantine (AD 1974).
Dibrachoides druso (Walker), from Europe, imported in unspecified year by unspecified
person, released against Hypera postica, current status unknown [Florida is not
listed among the states in which this insect was released in 1957-1975 by Dysart
& Day 1976] (Grant & Lambdin 1990). ~
Muscidifurax raptor Girault & Sanders, permit issued in 1971 for its release in Florida;
from Germany via France, imported in 1984 by P. B. Morgan as a biocontrol
agent for Musca domestic; from Hungary, imported in 1986-1987 by R. S. Pat-
terson; from Brazil, imported in 1987 and 1989 by P. B. Morgan and R. S. Patter-
son; from Hungary via France, imported in 1989 by D. R. Barnard; this species
was reported from Florida by Mitchell et al. (1974); the imported material was
used only for laboratory study, without release (Denmark & Porter 1973, AD
1984, 1987, 1989, P. B. Morgan pers. comm.).
Nasonia vitripennis (Walker), from France and from Zimbabwe, imported in 1986 by
P. B. Morgan against Musca domestic and Stomoxys calcitrans; from Brazil,
imported in 1987 by P. B. Morgan; this species may be native to Florida; the
imported material was used only for laboratory study, without release (AD 1986,
1987, P. B. Morgan pers. comm.).
Pachycrepoideus vindemiae (Rondani), from Mauritius, imported in 1974 and 1983;
from Brazil, imported in 1987 by P. B. Morgan; this species was reported from
Florida by Morgan & Patterson (1975) and was reported as attacking
Paratheresia claripalpis, a beneficial insect (see above), by Jaynes (1930); the
imported material was used only for laboratory study, without release (AD 1974,
1983, 1987, Stange 1986, P. B. Morgan pers. comm.).
Scutellista cyanea Motschulsky, from Italy, released in 1899 against Ceroplastes cir-
ripediformis, became established, attacking not only the target but two other
pests, Ceroplastes floridensis Comstock and Saissetia nigra (Nietner) (Bartlett
1978, Mizell 1990). *
Spalangia cameroni Perkins, from Mauritius and from Spain via France, imported in
1983 by P. B. Morgan against Musca domestic and Stomoxys calcitrans; from
France and from Germany via France and from Spain, imported in 1984 by P.
B. Morgan; from Zimbabwe, imported in 1986 by R. S. Patterson; from France,


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy 31

imported in 1987 by R. S. Patterson; from Morocco via France, imported in 1988
by R. S. Patterson; from Mauritius and from Brazil, imported in 1989; this species
was reported from Florida by Mitchell et al. (1974); the imported material was
used only for laboratory study, without release (AD 1983, 1984, 1986, 1987, 1989,
Stange 1986, P. B. Morgan pers. comm.).
Spalangia endius Walker, permit issued in 1971 for its importation and release in
Florida; from Thailand, imported in 1984 against Musca domestic and Stomoxys
calcitrans; from Mauritius via France and from Australia and from Hungary,
imported in 1986; from France and from India via France and from Brazil, im-
ported in 1987 by R. S. Patterson and P. B. Morgan; from Brazil, imported in
1989; this species was reported from Florida by Mitchell et al. (1974); the im-
ported material was used only for laboratory study, without release (Denmark
& Porter 1973, AD 1984, 1986, 1987, 1989, P. B. Morgan pers. comm.).
Spalangia gemina Bou6ek, from Brazil, imported in 1987 and 1989 by P. B. Morgan
against Musca domestic, not released (AD 1987, 1989, P. B. Morgan pers.
comm.).
Spalangia nigra Latreille, from France, imported in 1987 and 1990 by R. S. Patterson
against Musca domestic; this species was reported from Florida by Mitchell et
al. (1974); the imported material was used only for laboratory study, without
release (AD 1987, 1990, P. B. Morgan pers. comm.).
Spalangia nigroaenea Curtis, from Zimbabwe, imported in 1986 by R. S. Patterson
against Musca domestic and Stomoxys calcitrans; this species was reported
from Florida by Morgan & Patterson (1975); the imported material was used only
for laboratory study, without release (AD 1986, P. B. Morgan pers. comm.).
Spalangia sp., from Mauritius, imported in 1974 and 1983 by P. B. Morgan, not released
(AD 1974, 1983, P. B. Morgan pers. comm.).

HYMENOPTERA: SCELIONIDAE
Telenomus alecto (Crawford), from Trinidad via Louisiana, 2,200 adults released against
Diatraea saccharalis in 1950 at four sites, but no recoveries were made (Gifford
1964, Charpentier 1972, Bennett et al. 1990). ~
Telenomus remus Nixon, from Sarawak via Trinidad, imported in 1974-1975 by W. H.
Whitcomb against Spodopterafrugiperda, released in 1975-1977 in Dade County
(AD 1974, 1975, Wojcik et al. 1976, Grissell 1978, Waddill & Whitcomb 1980);
from the Cayman Islands, imported by F. D. Bennett in 1987, released in Dade
County in 1988-1989, without evidence of establishment; from Puerto Rico, im-
ported in 1987-1988 and 1990 by F. D. Bennett; from Jamaica, imported in 1989
by F. D. Bennett, but not released (AD 1987, 1988, 1989, 1990, Bennett & Capin-
era in press). ~
Telenomus sp., from Colombia, imported in 1986 by R. M. Baranowski and F. D.
Bennett (erroneously under the name T. alsophiliae Viereck) against Epimecis
detexta; 330 adults were shipped from quarantine to Homestead (Dade County),
but no releases were made (AD 1986, Mizell & Tedders 1990, F. D. Bennett pers.
comm., H. B. Glenn pers. comm.).
Trissolcus basalis (Wollaston), from Montserrat via Delaware, imported in 1973 by W.
H. Whitcomb and N. R. Spencer as a biocontrol agent for Nezara viridula,
though occurred already in Florida; from Australia, imported in 1974 by N. R.
Spencer (AD 1973, 1974, BIRL 1992, F. D. Bennett pers. comm.).
HYMENOPTERA: SERPHIDAE
Nothoserphus affisae (Watanabe), from Korea via Delaware, imported in 1990 into
quarantine as a potential biocontrol agent for Epilachna varivestis, but not re-
leased (BIRL 1990, Nong & Bennett in press).









Florida Entomologist 76(1)


TABLE 2. (Continued)

HYMENOPTERA: SIGNIPHORIDAE
Signiphora sp., from Brazil, imported in 1990 by F. D. Bennett together with
parasitoids of Bemisia tabaci, but probably is a hyperparasitoid and was termi-
nated in quarantine (AD 1990).
HYMENOPTERA: SPHECIDAE
Larra analis F., from Louisiana, imported by J. R. Watson in 1941-1942 against Scap-
teriscus spp., though it is a specific natural enemy of Neocurtilla hexadactyla
(Perty) which does not attack Scapteriscus, and though it occurred already in
Florida; all but one specimen were dead on arrival (Watson & Thompson 1943).
Larra bicolor F., from Para, Brazil via Puerto Rico, imported by R. I. Sailer, J. L.
Castner, and J. A. Reinert in 1981-1983 against Scapteriscus vicinus, S. borellii,
and S. abbreviatus, screened in quarantine and released in 1981 in Alachua,
Broward, and Hillsborough counties, in 1982 in Alachua, Broward, and Manatee
counties, in 1983 in Hillsborough County (AD 1981, 1982, 1983, ROBO 1981,
1982, 1983, Stange 1982), established only in Broward County and there studied
by J. L. Castner (Castner 1988), still (1992) with a very localized population in
Broward County; from Bolivia, imported [almost certainly together with material
of L. praedatrix] by F. D. Bennett in 1986-1989 and released in 1988-1989 in
Alachua County, without evidence of establishment (AD 1986, 1987, 1988, 1989,
F. D. Bennett pers. comm., Frank 1990). *
Larra godmani Cameron (under the name L. braunsii), from Santa Cruz, Bolivia,
imported and released against Scapteriscus spp. by F. D. Bennett in 1988-1989
in Alachua County, without evidence of establishment (Frank 1990, F. D. Ben-
nett pers. comm.). ~
Larra praedatrix (Strand), see under L. bicolor.
Larra spp., unidentified or under the names L. gastrica (Taschenberg) [synonym of L.
bicolor] and L. burmeisterii (Lynch) [the correct name is L. burmeisterii
(Holmberg)], from Uruguay, Bolivia, and Brazil, imported in 1984 and 1986 by
R. I. Sailer against Scapteriscus mole crickets; from Brazil, imported by F. D.
Bennett in 1989; all died in quarantine (AD 1984, 1986, 1989).
HYMENOPTERA: TORYMIDAE
Monodontomerus dentipes (Dalman), from Europe via North Carolina, imported and
released in 1969 by R. C. Wilkinson against Neodiprion lecontei, not established
(Wilkinson 1969, R. C. Wilkinson pers. comm.).
HYMENOPTERA: TRICHOGRAMMATIDAE
Brachyufens osborni (Dozier), from Puerto Rico, imported in 1969 (together with Test-
rastichus haitiensis) against Diaprepes abbreviatus, but no attempt was made
to culture it because it already was present in Florida as a parasitoid of
Pachnaeus; imported in 1977 but died during shipping (Sutton et al. 1972, AD
1977, Woodruff 1981).
Trichogramma evanescens Westwood, permit issued in 1968 for its importation and
release (Denmark & Porter 1973).
Trichogramma minutum Riley, permit issued in 1971 for its importation (Denmark &
Porter 1973).
Trichogramma plattneri Nagarkatti, from California, imported in 1985, reared on
Trichoplusia ni, 20,000 released in 1985 in Dade County against Epimecis de-
texta, but not established (Mizell & Tedders 1990, H. B. Glenn pers. comm.). ~
Trichogramma pretiosum Riley, from Texas, released inundatively in 1976 in Gadsden
County against Helicoverpa zea and Trichoplusia ni, established (Denmark &
Porter 1973, Martin et al. 1976, Jansson & Pefia 1990). *


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy 33

LEPIDOPTERA: COLEOPHORIDAE
Coleophora sp., from Brazil, imported in 1989 by D. H. Habeck against Schinus
terebinthifolius, but not reared and not released (AD 1989, D. H. Habeck pers.
comm.).
LEPIDOPTERA: NOCTUIDAE
Spodoptera pectinicornis (Hampson), from Thailand, imported in 1986-1988 by D. H.
Habeck and C. R. Thompson against Pistia stratiotes, released in 1990 in Glades,
Palm Beach and St. Lucie counties, and in 1991 in Brevard, Broward, Gadsden,
Glades, Okeechobee, Putnam, and Sumter counties; it is not yet clear that estab-
lishment is permanent (AD 1986, 1987, 1988, Buckingham & Habeck 1990, Dray
& Center 1992, D. H. Habeck pers. comm.). ?
LEPIDOPTERA: PYRALIDAE
Acentria ephemerella (Denis & Schiffermuller), native to the northern USA, from New
York via Maryland, imported in 1975-1976 (under the name Acentropus niveus)
by N. R. Spencer, and in 1978 by G. R. Buckingham, against Myriophyllum
spicatum, but died in quarantine (AD 1975, 1976, 1978, G. R. Buckingham pers.
comm.).
Acigona infusella (Walker), from Argentina, imported in 1974-1975 by N. R. Spencer
against Eichhornia crassipes, but was not released, and died in quarantine (AD
1974, 1975, N. R. Spencer pers. comm.).
Parapoynx diminutalis Snellen, from Asia via Panama, imported in 1980-1982 by G.
R. Buckingham against Hydrilla verticillata, though by then was established in
Florida as an immigrant (AD 1980, 1981, 1982, G. R. Buckingham pers. comm.).
Parapoynx stratiotata L., from Italy via Delaware, from Yugoslavia, and from Italy,
imported in 1975-1976 by N. R. Spencer as a potential biocontrol agent for
Myriophyllum spicatum, but was terminated in quarantine (AD 1975, 1976,
BIRL 1992, D. H. Habeck pers. comm., N. R. Spencer pers. comm.).
Sameodes albiguttalis (Warren), from Argentina, imported in 1975-1976 by N. R.
Spencer, released in Broward, Collier Dade, and Pinellas counties in 1977-1979
(numbers not counted) and in 1979-1980 in Alachua County (79,093 in 1979, 19,764
in 1980) against Eichhornia crassipes, established by 1979 (AD 1975, 1976,
Center & Durden 1981, Center 1984, Buckingham & Habeck 1990). *
Vogtia malloi Pastrana, from Argentina, released in 1971-1972 in Alachua, Broward,
Duval, Orange and St. Lucie counties against Alternanthera philoxeroides, es-
tablished (Brown & Spencer 1973, Coulson 1977, Buckingham & Habeck 1990). *
LEPIDOPTERA: TORTRICIDAE
Episimus utilis Zimmerman, from Brazil, imported in 1989 and 1991 by D. H. Habeck
against Schinus terebinthifolius, but not released (AD 1989, 1991, D. H. Habeck
pers. comm.).
NEUROPTERA: CHRYSOPIDAE
Chrysoperla carnea (Stephens), from India, 8,500 eggs were released in 1957 in five
fields, and 1,150 were released in 1958 in five fields (from Belle Glade, Palm
Beach County) against Sipha flava, and others were released from Quiney
(Gadsden County) against Myzus persicae and Therioaphis maculata; not estab-
lished (Denmark 1964, Charpentier et al. 1972, Jackson 1990, L. A. Stange pers.
comm.). ~
STREPSIPTERA: STYLOPIDAE
Stenocranophilus quadratus Pierce, from Jamaica via Trinidad and New Jersey, 580
adults and nymphs were released in 1959 at one site from Belle Glade (Palm
Beach County) against Saccharosydne saccharivora, established (Bennett 1960,









34 Florida Entomologist 76(1) March, 1993

TABLE 2. (Continued)

Denmark 1964, Gifford 1964, Simmonds 1969, Charpentier et al. 1972, Bennett
et al. 1990, F. D. Bennett pers. comm.). *
THYSANOPTERA: PALEOTHRIPIDAE
Amynothrips andersoni O'Neil, from Argentina, released in 1967-1972 in Alachua, Bro-
ward, Clay, Duval, Glades, Orange and Palm Beach counties against Alternanth-
era philoxeroides, established (Coulson 1977, Buckingham & Habeck 1990). *
Liothrips ichinii Hood, from Brazil, imported into quarantine in 1989-1991 by F. D.
Bennett and D. H. Habeck against Schinus terebinthifolius, but no releases were
made (AD 1989, 1990, 1991, Habeck et al. in press, F. D. Bennett pers. comm.)


became established and caused severe problems. Control was first attempted by phys-
ical methods, which already had been shown to be ineffective in California. In 1899, a
new attempt was made to import the biocontrol agent, resulting in completely successful
control. A second introduced insect species, Gromphadorina sp. (Blattaria: Oxyhyd-
roidae), is a potential pest, but is not yet known to be established (see Table 2). The
other introduced pests are plants that have become weeds. They were brought to
Florida either as ornamentals [Eichhornia crassipes (Martius) Solms (waterhyacinth),
Melaleuca quinquenervia (Cavanilles) S. T. Blake melaleucaa), Schinus terebinthifolius
Raddi (Brazilian peppertree)], or by the aquarium trade [Hydrilla verticillata (Lf.)
Royle (hydrilla), Myriophyllum spicatum L. (Eurasian watermilfoil)], and then became
feral and difficult to control.
A problem arises with some pests that have been widely distributed for such a long
time that their native ranges are not clear to taxonomists. An example among the
insects is Selenothrips rubrocinctus (Giard) (Thysanoptera: Thripidae), which is widely
distributed in the tropics and subtropics around the world. An example among the
plants is the aquatic weed Pistia stratiotes L. (waterlettuce), which was observed in
Florida in the 18th century but yet may not be native. In a few instances, particularly
where a specialist pest of an introduced plant is concerned, the pest has been assumed
to be an immigrant, from the same part of the world as its host plant; an example is
Chrysomphalus aonidum (L.) (Homoptera: Diaspididae), whichis assumed to be native
to Asia, and whose common name "Florida red scale" is thus a misnomer. In other
instances, when all the closest relatives of a pest are from one continent, it has been
assumed that the pest likewise is from that continent; an example is Aedes aegypti (L.)
(Diptera: Culicidae), whose closest relatives are African. Counter-examples to both of
these assumptions can be found; therefore, we must view any conclusions drawn from
the assumptions as tentative.

Introduced Insects

We included information, when we had it, on geographical origins, dates of introduc-
tion, locations of releases, and the status of each introduced insect (Table 2). The latter
kind of information distinguishes among species imported but not released, species
released but not established, species released and established, and species whose status
is currently unknown. Establishment (see Hall & Ehler 1979, Stiling 1990) was
documented by record of subsequent recovery in the field, although this criterion may
yield questionable conclusions, for at least three reasons. First, little or no follow-up
work has been done for many of the releases. Second, species that establish in the short
term may fail to do so in the long term, and species that seem to fail to establish in the
short term may eventually do so (a temporal scale problem, which would be reduced if









Insect Behavioral Ecology-'92: Frank & McCoy


an appropriate time frame were specified in advance). Third, establishment may be
judged to have occurred in one part of the state but not in another (a spatial scale
problem, which would be reduced if Florida were not considered the geographical area
of interest). We also included additional information, such as on sizes of releases, that
we had for some species. Because we did not feel we possessed enough information to
do so, we made no particular attempt to draw conclusions about the success (see Hall
et al. 1980, Stiling 1993) of releases for biological control in Florida. The cut-off date is
1991 for new importations.
Almost all of the introduced insects that we included in our tabulation were imported
deliberately and were deemed beneficial and suitable for release (but see Howarth
1991). Release of the biocontrol agents usually was subject to proof of host-specificity,
however. Almost all also were not already established in Florida, although in a few
instances, strains (subspecific or infra-subspecific categories) from other areas were
imported. We included on the list: (1) a few species shipped unsolicited to the quarantine
facility of the Florida Biological Control Laboratory by foreign suppliers, but which
were terminated in quarantine, (2) a few unexpected parasitoid species that emerged
from their imported hosts in the quarantine facility, and (3) two species imported by
members of the public without permit and which are documented to have been released
in Florida. We omitted from the list: (4) any pest species imported into the quarantine
facility simply as a host for beneficial parasitoids or predators, and which was termi-
nated in the quarantine facility, (5) species imported by the pet trade or by members
of the public without permit or other public record, (6) species imported under permit
for experimental purposes in secure laboratories, (7) species imported under permit for
educational purposes but which were not intended for release (such as exotic but-
terflies), and (8) beneficial species imported under permit by commercial organizations
as biological control agents for pests. Records of the permits for categories (6)-(8) exist,
but records of receipt of shipments for categories (7)-(8) are not in the public record.
The last category (8) is of insect biocontrol agents produced in insectaries in other
states, imported into Florida under permit as biopesticides by commercial companies,
but not passed through quarantine; no public record is made of their numbers or places
of release, and no special record is made of those among them which are not established
already in Florida; such importations could lead to establishment of species which are
not already present in Florida. Icerya purchase is not included in out table of importa-
tions because it is not a biological control agent and because we discuss its introduction
above.
Alternative scientific names have been used in the literature for some of the biolog-
ical control agents listed in Table 2. Table 3 shows these alternative names, and will
help anyone who wants to relate information from Table 2 to the literature.

RESULTS AND DISCUSSION

Targets of Classical Biological Control Efforts

A diverse group of 75 organisms has been targeted for classical biological in Florida
(Table 1). Slightly less than half (48%) of the insects belong to the order Homoptera.
Other well-represented insect orders are Lepidoptera (24%) and Coleoptera (10%). Two
families, Diaspididae and Aphididae, together include half of the species of Homoptera.
Single families, Noctuidae and Curculionidae, respectively include nearly half (44%) of
the species of Lepidoptera and more than half (57%) of the species of Coleoptera. The
distribution of targeted insect pest species among orders is not very similar to the
distribution of recent immigrant insect species among orders (see Frank & McCoy
1992). The eight targeted plant pest species are from eight orders.
Most (79% of the insects, 75% of the plants) of the pests targeted for classical









36 Florida Entomologist 76(1) March, 1993

TABLE 3.Alternative scientific names have been used in the literature for several of
the biological control agents listed in these pages. Synonyms were-used inad-
vertently in many instances, but in some instances misspellings were used,
or misidentifications were made. The list following shows (at left) names that
have been used in the literature, and (at right) names that we believe to be
correct now for the species in question.

for Acentropus niveus see Acentria ephemerella
for Adenis variegata see Hippodamia variegata
for Adonia variegata see Hippodamia variegata
for Aganaspis daci see Trybliographa daci
for Agathis stigmatera (Cresson) see Alabagrus stigma
for Apanteles flavipes see Cotesia flavipes
for Aphelinus semiflavus (Howard) see Aphelinus asychis
for Aphis citricola van der Goot see Aphis spiraecola
for Aphycus helvolus see Metaphycus helvolus
for Aphycus luteolus see Metaphycus luteolus
for Aphytis citrinus (Compere) see Aphytis aonidiae
for Aptesis basizonia see Pleolophus basizonus
for Aspidiotiphagus see Encarsia
for Asynonychus godmani see Asynonychus godmanni
for Azya trinitatis see Pseudoazya trinitatis
for Bagous pulchellus Hustache see Bagous affinis
for Bassus stigmaterus (Cresson) see Alabagrus stigma
for Biosteres longicaudatus see Diachasmimorpha longicaudata
for Biosteres oophilus (Fullaway) see Biosteres arisanus
for Bracanastrepha anastrephae see Utetes anastrephae
for Bracon brevicornis see Habrobracon brevicornis
for Bracon chinensis see Myosoma chinensis
for Bracon vesticida see Bracon vestiticida
for Brumus suturalis see Brumoides suturalis
for Centeterus alternecoloratus see Aubertillus alternecoloratus
for Ceromasia sphenophori see Lixophaga sphenophori
for Chilomenes sexmaculata see Menochilus sexmaculatus
for Chrysopa carnea see Chrysoperla carnea
for Coccinella punctata see Coccinella septempumctata
for Cryptochaetum see Cryptochetum
for Cryptochetum monophlebi Skuse see C. iceryae
for Cryptognatha flavescens see Serangium flavescens
for Cryptognatha simillima Sicard see Cryptognatha gemellata
for Dasyscapus parvipennis Gahan see Goetheana parvipennis
for Delphastus sonoricus Casey see Delphastus pusillus
for Dibrachoides dynastes (Foerster) see Dibrachoides druso
for Doryctobracon cereum (Gahan) see Doryctobracon areolatus
for Dusmetia sangwani see Neodusmetia sangwani
for Encarsia tabacivora Viggiani see Encarsia pergandiella
for Eocanthecona furcellata see Cantheconidia furcellata
for Euphasiopteryx depleta see Ormia depleta
for Goetheana parvipennis (Gahan) see Goetheana shakespearei
for Gonatopus sp. see Pseudogonatopus variistriatus
for Ipobracon rimac see Iphiaulax rimac
for Heterolaccus grandis see Catolaccus grandis
for Iphiaulax amabilis see Digonogastra amabilis









Insect Behavioral Ecology-'92: Frank & McCoy 37

for Iphiaulax rimac see Digonogastra rimac
for Larra braunsii Kohl see Larra godmani
for Leis dimidiata see Harmonia dimidiata
for Leptomastix abnormis seee Leptomastidea abnormis
for Litodactylus leucogaster see Phytobius leucogaster
for Litomastix truncatella (Dalman) see Copidosomafloridanum
for Lydella grisescens Robineau-Desvoidy see Lydella thompsoni
for Microbracon vestiticida see Bracon vestiticida
for Microcharops bimaculata (Ashm.) see Microcharops anticarsiae
for Microceromasia sphenophori see Lixophaga sphenophori
for Namangana pectinicornis see Spodoptera pectinicornis
for Nephaspis amnicola Wingo see Nephaspis oculata
for Opius anastrephae see Utetes anastrephae
for Opius concolor see Psyttalia concolor
for Opius fletcheri see Psyttalia fletcheri
for Opius incisi see Psyttalia incisi
for Opius oophilus Fullaway see Biosteres arisanus
for Opius persulcatus Silvestri see Biosteres vandenboschi
for Opius trinidadensis see Doryctobracon trinidadensis
for Opius tryoni see Diachasmimorpha tryoni
for Opius vandenboschi see Biosteres vandenboschi
for Pantomorus cervinus (Boheman) see Asynonychus godmanni
for Pantomorus godmanni see Asynonychus godmanni
for Parachasma cereum (Gahan) see Doryctobracon areolatus
for Parachasma crawfordi see Doryctobracon crawfordi
for Parachasma trinidadense see Doryctobracon trinidadense
for Paraleptomastix abnormis see Leptomastidea abnormis
for Phania vittata see Evbrissa vittata
for Pimpla sp. see Coccygomimus sp.
for Praon palitans Muesebeck see Praon exsoletum
for Prospaltella see Encarsia
for Pseudoarchytopsis see Archytas
for Pseudococcobius terry Fullaway see Pseudaphycus mundus
for Pyrophorus luminosus see Ignelater luminosus
for Scapteriscus acletus Rehn & Hebard see Scapteriscus borellii
for Scymus binaeratus see Nephus binaevatus
for Stethorus atomus Casey see Stethorus utilis
for Theresia claripalpis see Paratheresia claripalpis
for Timberlakia europaea see Pseudectroma europaea
for Triaspis vesticida see Triaspis vestiticida
for Trioxys utilis Muesebeck see Trioxys complanatus



biological control in Florida are known to be adventive. Although the adventive insect
pests had their geographic origins in many regions of the globe, a large percentage
(43%) is native to Asia, and most of these Asian natives attack citrus. These Asian pests
are atypical of adventive insects in 1971-present, because most of the latter are of
Neotropical origin (Frank & McCoy 1992). However, many of the targeted insect pests
were pests of citrus, which is of Asian origin. The targeted adventive plant pests are
from South America (50%), Eurasia (33%), and Australia (17%). The early part of the
pattern of arrival of targeted pest species in Florida (Fig. 1) may illustrate the linkage
of pest exchange with commercial traffic noted by Sailer (1983). The importation of









Florida Entomologist 76(1)


<18501850186018701880189019001910 19201930194019501960197019801990

DECADE

Fig. 1. Arrival of targeted pest species in Florida. Line is the cumulative number
of species. Note that one species, Aleurocanthus woglumi Ashby, immigrated to Florida
twice (Table 1).


several ornamental plants that later became pests near the end of the nineteenth
century further inflated the number of arrivals at that time.
It is interesting to note that acting under the Plant Pest Act of 1912 and Plant
Quarantine Act of 1957, USDA agricultural inspectors have for decades tried to exclude
phytophagous insects ("plant pests") from entry into the USA, yet the USDA has
encouraged, and itself has taken part in, importation of exotic plants as ornamentals.
This incongruity is explicable in terms of trade: sales of exotic terrestrial plants (by the
nursery trade), exotic aquatic plants (by the aquarium trade), and exotic animals, espec-
ially vertebrates, but also mollusks and anthropods (by the pet trade) provide a profit
to importers. Our laws make it acceptable to import potential pests if a profit is to be
made, but not to import worthless (i.e., unsalable) potential pests. We are not aware
of anything in the laws that requires importers to pay for the control of imported
organisms that have become pests, nor even to pay the cost of research toward finding
means of control of such pests, although it strikes us as fair that they should do so.
None of the targeted pests known to be native (16% of the insects, 12% of the plants)
is precinctive (i.e., occurring only in Florida; see Frank & McCoy (1990) for definition
of this term). All have wider distributions, either in the southeastern states generally,
or also in the West Indies, with Florida only a part of their native ranges. An example
among the insects is Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae),
which has a wide distribution in the West Indies, Central and South America; it occurs
in the southern USA, but in severe winters it can survive only in the extreme south of


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy 39

Florida. An example among the plants is Lantana camera L. (lantana), which occurs
naturally from the southern USA to northern South America.

Introduced Insects

Our tabulation of insects introduced into Florida (Table 2) includes 351 taxa (a few
species are lumped together). Of these 351 taxa, 32 appear to have occurred already in
Florida; 24 apparently are natives and 8 apparently are immigrants. Some of the immi-
grants, such as Bathyplectes curculionis (Thomson) (Hymenoptera: Ichneumonidae),
seem to have arrived in Florida by immigration from other states into which they had
been imported, while others, such as Megastigmus transvaalensis (Hussey) (Hymenopt-
era: Torymidae) and Utetes anastrephae (Viereck) (Hymenoptera: Braconidae), may
have arrived directly from abroad by stowing away in cargoes of plants and other mater-
ials, probably with their host insects (Frank & McCoy 1992). Another 11 taxa are
thought to have been introduced because permits were issued for their importation, but
no actual records of importation could be located. Exactly half, 154, of the remaining
308 taxa were released in Florida. Three of the species released are not biological
control agents, leaving 151 taxa, of which 139 targeted insect pests and 12 targeted
plant pests. Biological control agents have been released in Florida since the 1890s (Fig.
2).
Overall, the proportion of establishment of biological control agents in Florida is
27.8%. If the agents targeting insect and plant pests are considered separately, the


<1850185018601870188018901900191019201930194019501960197019801990

DECADE
Fig. 2. Release of insect biological control agents in Florida (one release = one agent
taxon introduced against one pest taxon in one decade). Upper line is the cumulative
number of releases, lower line is the cumulative number of releases that resulted in
establishment.









40 Florida Entomologist 76(1) March, 1993

proportions are 24.5% and 66.7%, respectively. If the taxa whose current status is
unknown are assumed to establish in the same proportions as the taxa whose status is
known, then the overall proportions increase to 27.3% and 91.7%, respectively. Our
proportion of establishment for agents targeting insect pests, calculated in either way,
loosely matches what Hall & Ehler (1979) refer to as the "often quoted figure of 0.20-
0.25" (see Clausen 1956). Our proportion of establishment, however, is less than the
worldwide proportions calculated from published lists by Hall & Ehler (1979) (34%) and
Stilirg (1990) (35.6% or 43.4%, depending upon how the calculations were made).
It may be significant that the establishment proportion calculated by Hall & Ehler
(1979) for California (22%) also seems to be relatively low. They attributed the relatively
low proportion in California to that state's efficient documentation of introductions and
its role as a "screening ground"; for natural enemies, both of which would result in
proportionately more failures being reported. To a certain degree, the same two expla-
nations hold for Florida.
Proportions of establishment may differ in some systematic way among species or
among situations, but the factors that correlate with relatively high or relatively low
proportions of establishment of biological control agents seem to be complex and poorly
understood (Gross & Pair 1986, Stiling 1990; see also Murdoch et al. 1985). Few research-
ers today would try to predict which species-introductions are likely to succeed, as
was common practice at one time (see DeBach 1971, Huffaker et al. 1971, Price 1972).
Previous studies have reached several relevant conclusions concerning proportions of
establishment of insect agents, however. (These same conclusions often apply equally
well to proportions of success.) Proportions are similar for predators and parasitoids
(Hall & Ehler 1979); proportions are higher (1) against certain orders of pest than
against others (Hall & Ehler 1979, Stiling 1990), (2) against exotic pests than against
native ones (Hall & Ehler 1979, Stiling 1990), (3) against exposed pests than against
concealed ones (Stiling 1990, Gross 1991; see also Hawkins & Lawton 1987), (4) against
monophagous pests than against polyphagous ones (Martin et al. 1981, Stiling 1990), (5)
in "stable" habitats than in "unstable" ones (Hall & Ehler 1979, but see Stiling 1990),
(6) when agents are adapted to the climatic regimes in which the pests reside than when
they are not (Messenger 1971, Stiling 1990), (7) when large numbers of individuals of
agents are introduced than when they are not (Beirne 1975, but see Gross & Pair 1986).
Proportions of introduced species becoming established in the USA declined over time
(Hall & Ehler 1979). Some factors that may interfere with establishment, but that we
shall not discuss, are voltinism (van den Bosch & Messenger 1973, Stiling 1990), lack
of alternate prey/hosts (van den Bosch & Messenger 1973, Eikenbary & Rogers 1974),
lack of foodstuff for adult parasitoids (Hagen et al. 1970, van den Bosch & Messenger
1973), and most controversially competition (Ehler & Hall 1982, 1984, Keller 1984,
Simberloff 1986, Myers et al. 1989; see example in Schuster & Dean 1976 and Anagyrus
antoninae Timberlake in Table 2).
Examination of these conclusions with our data set led to some interesting results.
The proportions of establishment of insect predators and parasitoids are 26.7% and
23.9%, respectively. These proportions cannot be shown to be different (G-statistic =
0.04, P > 0.05), but both are lower than the proportion of establishment of herbivores
(66.7%) (G-statistics = 88.02 and 12.34, P < 0.05). Most releases were against targets
in two orders, Homoptera and Lepidoptera, and the proportions of establishment are
different for them: 31.7% and 10.5%, respectively (G-statistic = 6.82, P < 0.05). (Num-
bers of releases against targets in other orders were too few to draw meaningful conclu-
sions.) Although the worldwide proportions of establishment they report for these two
orders are substantially higher than ours, Hall & Ehler (1979) and Stiling (1990) both
conclude that the proportion of establishment for homopterous targets exceeds the
proportion for lepidopterous targets.









Insect Behavioral Ecology-'92: Frank & McCoy 41

We could not examine four of the conclusions with our data set. Because most of the
native insect pests are in the order Lepidoptera, and we have just shown pests in this
order to be relatively poor candidates for establishment, we could not compare propor-
tions of establishment against native and exotic pests. Likewise, pests in the order
Homoptera, which we have shown to be relatively good candidates for establishment,
tend to occur in large groups on the surface of one or a few plant taxa, particularly on
citrus trees; therefore, because pests in this order make up nearly half of all the targeted
insects, we also could not compare proportions of establishment against exposed and
concealed pests, against monophagous and polyphagous pests, or in "stable" and unsta-
ble" habitats.
We examined the effect of climatic regime in the useful, albeit indirect, way
suggested by Stiling (1990). It is necessary to employ an indirect method because direct
measures of local climate usually are not available for either the pest's location or the
biological control agent's origin (see Stiling 1990). Because Stiling's (1990) method oper-
ates on a relatively large spatial scale, any results derived from its use must be consid-
ered tentative. Stiling's (1990) results suggest that, for Florida, which we will consider
mostly temperate, agents of temperate origin will be more readily established than
agents of tropical origin. We could not confirm this tendency for agents targeting insect
pests; the establishment proportions are 22.2% and 29.0% for agents probably of temp-
erate and tropical origin, respectively (G-statistic = 0.48, P > 0.05). When the regions
of origin are grouped as New World, Eurasia, tropical Asia/Australia, Africa, and
Hawaii, the resulting establishment proportions range from 20.0% to 33.3%, none of
which can be shown to be different from the others (G-statistics = 0.00 to 0.08, P >
0.05). Proportions of establishment of agents of island origin (24.2%) are not different
from those of agents of mainland origin (27.1%) (G-statistic = 0.06, P > 0.05). Agents
from India/Pakistan and the West Indies have the highest (36.0%) and lowest (17.6%)
proportion of establishment, respectively, for individual regions, but neither proportion
can be shown to be different from that of the rest of the globe (G-statistics = 1.42 and
0.64, P > 0.05).
Although agents from specific regions cannot be shown to be unusual in proportion
of establishment, more detailed examination of the cases of India/Pakistan and the West
Indies may reveal patterns that are potentially of interest. Most of the agents acquired
from India/Pakistan targeted insects in the order Homoptera, and many of the ones
that established targeted more than one pest. These facts may help to explain their
relatively high proportion of establishment. The relatively low proportion of establish-
ment for agents acquired from the West Indies is less easily explained, especially as
immigration from the West Indies appears to occur regularly (Frank & McCoy 1992).
Many of these agents also targeted insects in the order Homoptera, but the lists of
targets of agents from India/Pakistan and from the West Indies overlap very little.
Perhaps the particular pests targeted by West Indian agents simply are poorer candi-
dates for establishment than those targeted by agents from other regions; but, of course,
such a conclusion begs the question.
The mean number of individuals released for agents that established was quite near
the mean number released for agents that failed to establish. The mean numbers were
4743 (7364) individuals and 4332 (10,979) individuals, respectively, when releases
against Anastrepha suspense (Loew) (Dip.: Tephritidae) are excluded. Inclusion of re-
leases of A. suspense, one of which exceeds one million individuals, raises the mean
number of individuals released for agents that established to 30,377 (161,726). The
variances are too high to allow meaningful conclusions to be drawn, but even if they
were not, we are not sure what a difference in numbers of individuals might mean. If
number of individuals released could be shown to be related positively to chance of
establishment, the result might mean that raising extremely large numbers of individ-










Florida Entomologist 76(1)


uals for release could improve the chance of establishment. Release of large numbers
of individuals in more locations, targeting more microclimates, could improve chances
of success. On the other hand, the result could simply mean that large numbers of
individuals are released only when the chance of establishment (and, perhaps, success)
is thought to be high in advance. It seems likely to us that releases of large numbers
of individuals are warranted sometimes, but that at present we have little way of
predicting whether release of large numbers is justifiable.
No decline in proportion of establishment over time was obvious. We did discern an
apparent tendency for proportions to be lower for decades in which relatively many
releases were made, which we believe indicates that occasional "shotgun"; releases are
made, and that many of these releases have relatively little chance of establishment.
The tendency indeed is present, but it is not a particularly strong one (Spearman's r =
-0.515, P > 0.05).
We also looked for patterns in proportions of establishment relative to location of
release. Releases were known to have been made in 41 of the 67 counties, and the
outcome was known for at least one release in 39 of these counties. Only five counties
had proportions of establishment less than 50%: Alachua, Dade, Gadsden, Indian River,
and Palm Beach. Coincidentally, four of the five (all but Indian River) were the sites
of the largest numbers of releases. We refer to these counties as "donors," where
potential agents are released and screened; agents judged to have a high chance of
establishment are then released in other counties, which we refer to as "recipients."
Because proportions of establishment were greater in Dade (37.0%), Indian River
(37.5%), and Palm Beach (41.4%) counties than in Alachua (12.5%) or Gadsden (7.7%)
counties, we thought that we might also find greater proportions of establishment in
recipient counties in the south or along the coast than in recipient counties in the north
or inland. No such correlations were obvious to us, however, but trends might be
masked if agents thought in advance to have a high chance of establishment were
released preferentially in recipient counties. The generally low proportions of success
in the Panhandle region may merit investigation, but sample sizes are too small for us
to reach any firm conclusions.
What needs to be done to further our understanding of how insects introduced into
Florida react to the environment that they encounter? An analysis of proportions of
success is needed, to accompany the analysis of proportions of establishment. Some
greater insight into the behavioral ecologies of biological control agents and their targets
also is needed. We suggest that closer scrutiny of all of the introductions, including the
biologies and ecologies of the organisms involved and, perhaps, the use of multiple
comparisons, would prove productive. On a larger scale, in order to determine how
Florida fares relative to other regions, two other pieces of information would be valu-
able: (1) the variation in the way proportions of establishment and success are calculated
and reported worldwide and (2) a list of the taxa that were imported but not released
worldwide ("failed to establish or succeed in the laboratory"). The latter piece of infor-
mation might well alter the way in which we think about proportions of establishment
and success, although we have not considered all the implications.

THE SYMPOSIUM

In dealing with the behavioral ecology of immigrant insects as the theme of last
year's symposium, we faced two unknowns: we did not know how many individuals of
any immigrant species arrived in Florida, nor precisely how these immigrations were
distributed in time. This restricted us to dealing with successful immigrations, i.e.,
immigrant species which were shown to have established populations in Florida.
The theme of this year's symposium is introduced insects. People have introduced


March, 1993









Insect Behavioral Ecology-'92: Frank & McCoy


insects into Florida for various reasons, but very nearly all of the introductions that
have been documented were for biological control purposes. The Florida Entomological
Society had planned a separate symposium on biological control. When it became clear
that much of our symposium would be about the behavioral ecology of biological control,
we decided to merge the two symposia. Most of the contributions are about the be-
havioral ecology of insects or nematodes introduced for biological control purposes, but
some of the contributions are about related topics that we found interesting.
Practitioners of classical biological control import biological control agents against
immigrant pests and sometimes against native pests. These introductions are made
when native or previously-introduced biological control agents do not control the pests
adequately. Fred Bennett reviews reports of the effect of newly-introduced agents on
the previously-occurring natural enemies; he finds that populations of the latter some-
times are reduced, and that aggression of one biological control agent against another
has been documented.
Pest ants have been the targets of biological control attempts, especially using patho-
gens. David Oi and Roberto Pereira review the literature on the behavior of ants
afflicted by bacteria, fungi, protozoans, nematodes and trematodes. They document
production of secretions, grooming, nest hygiene, avoidance, and dispersal as behaviors
displayed by ants of a range of genera against such natural enemies.
Entomopathogenic nematodes have been used as biopesticides, typically are applied
at high dosages, and have no residual effect because they do not reproduce in the
habitats where they are applied. Patrick Parkman and Howard Frank describe a novel
use of an imported nematode species which reproduces in its specific host. They at-
tracted mole crickets to synthetic mole cricket song, there exposed the mole crickets
to the nematodes, and then permitted the insects to disperse. Thus, they fomented
disease epidemics in mole cricket populations on Florida golf courses.
The theme of use of entomopathogenic nematodes to control insects is continued by
Rick Jansson. He examines behavioral characteristics of heterorhabditid and steiner-
nematid nematodes collected in recent surveys of soils. Many of the species were previ-
ously unknown taxonomically and behaviorally. He is rightly concerned about their
host-range and about potential effects of releasing them in places where they are not
native.
Waterhyacinth is one of the most invasive aquatic weeds infesting waterways in the
southern USA. Two weevil species of the genus Neochetina have been introduced
against it, but chemical herbicides are still in use. Kim Haag (manuscript not submitted)
shows that the weevils can reduce waterhyacinth populations. When waterhyacinth in
a lake was treated experimentally with 2,4-D, weevils aggregated in untreated refugia
and there caused significant reduction in plant stature and growth.
Coccinella septempunctata, a widely-distributed palearctic ladybird, was released
first in the USA in 1956, and further releases were made in 1958-1973. Established
populations were found in 1973 in Canada and New Jersey. John Obrycki (manuscript
not submitted) examines behavioral characteristics of this insect in attempt to determine
why it took so long to become established. He also is concerned with documenting any
effects of this species, which is one of the least prey-specific of ladybirds, on the ladybird
fauna native to North America.
Under what circumstances is a classical biological control project justifiable econom-
ically? Michael Habeck (and two co-authors) are economists who look critically at project
cost, length, chance of success, and the time value of money to build a simple mathemat-
ical model. By their calculations, the benefits of a biological control project must be
predicted to exceed about $62,000 per year before its funding can be justified. Of course
there are exceptions, but biological control practitioners need to understand the
economic realities when they seek funding for research.










Florida Entomologist 76(1)


Gypsy moth was introduced into Massachusetts in the last century, escaped from
captivity, and became a pest. It has been the target of biological control by introduction
of natural enemies from the palearctic region, but these introductions were not reviewed
by Jon Allen (and eight co-authors). Instead, these colleagues address the probability
of gypsy moth managing to colonize Florida from northern states. They conclude that
the moth probably will establish populations in northern Florida, but its behavioral
response to mild winters and sparsely-distributed host-trees may not allow it to achieve
pest status.
ACKNOWLEDGMENTS

We thank Jack Coulson (USDA-ARS, Beltsville, MD) and Larry Ertle (USDA-ARS,
Newark, DE) for providing information from the databases that they maintain (ROBO
and BIRL respectively, see references cited), Avas Hamon for taxonomic information
on Homoptera, Frank Mead on Hemiptera, Lionel Stange on Neuroptera, Dale Habeck
and John Heppner on Lepidoptera, John Foltz on forest insects, Harold Denmark on
Aphididae and Thysanoptera, Charles O'Brien on Curculionidae, Virendra Gupta on
Ichneumonidae, and Robert Wharton on Braconidae, Robert Woodruff for help in track-
ing specimens, and the literature, on Coccinellidae and Scarabaeidae, Christine Bennett,
Harold Denmark, Avas Hamon, and Limhuot Nong for help locating records of permits
and importations, and the following for further information on the insects which they
and their colleagues imported: Fred Bennett, Harold Browning, Gary Buckingham,
John Capinera, Arnold Drooz, Holly Glenn, Dale Habeck, David-Hall, Rick Jansson,
Gary Leibee, John McLaughlin, Everett Mitchell, Philip Morgan, Ru Nguyen, Jorge
Pefia, Eric Schreiber, Omelio Sosa, Neal Spencer, Van Waddill, Willard Whitcomb,
Robert Wilkinson, and Dan Wojcik. Especial thanks to Fred Bennett for commenting
on errors and additions to the manuscript at several points in its development. Peter
Stiling and John Capinera kindly reviewed the manuscript. We thank Julio Arias for
translating the abstract into Spanish. We apologize to any contributor whose name we
have omitted. This is University of Florida, Institute of Food & Agricultural Sciences,
journal series no. R-02957.

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SIMMONDS, F. J. 1969. Biological control of sugar cane pests: A general survey, p.
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STANGE, L. A. 1982. Biological control laboratory. Florida Dept. Agric. Consumer
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STILING, P. 1990. Calculating the establishment rates of parasitoids in classical biolog-
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SUMMERS, T. E., E. G. KING, D. F. MARTIN, AND R. D. JACKSON. 1976. Biological
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SUTTON, E. A., A. G. SELHIME, AND W. MCCLOUD. 1972. Colonization and release
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54 Florida Entomologist 76(1) March, 1993

DO INTRODUCED PARASITOIDS DISPLACE NATIVE ONES?

FRED D. BENNETT
Entomology and Nematology Department
University of Florida
P.O.Box 110620
Gainesville, FL 32611-0620

ABSTRACT

An attempt was made to determine from the literature and from experience whether
native parasitoids of native pests are eliminated by introduced parasitoids. Several
native or previously-introduced parasitoids of exotic pest species have become scarce
or apparently have disappeared completely following the introduction of additional nat-
ural enemies. It is concluded that native parasitoids can be displaced over much of their
range, but theie usually are favorable habitats where they are able to co-exist with the
introduced species.

RESUME

Se quiere determinar en base a lo reportado en la literature y en base a experiencia
propia si los parasitoides natives de las plagas nativas son eliminados al introducir
parasitoides foraneos. Algunos parasitoides nativos o introducidos previamente contra
plagas exoticas se han reducido en numero o han desaparecido completamente despues
de la introduction de enemigos adicionales. Se concluye que los parasitoides nativos
pueden ser desplazados fuera de su rango, pero usualmente encuentran habitats favor-
ables donde ellos pueden co-existir con las species introducidas.




During the discussion period in one of the biological control symposia of the XIX
International Congress of Entomology held in Beijing, China, I inquired whether any-
one could cite an example in which an introduced parasitoid had displaced a native
parasitoid of a native pest to the point of extinction. Nobody could provide an example.
If there are instances where this occurred, the chances that they would be recorded are
very slim, mainly because of the frequent absence of intensive pre-introduction studies
of the parasitic fauna. Hence, later, if only introduced parasitoids were reared from a
native pest, we could not be certain whether they in fact had displaced native parasitoids
or whether there had been any native parasitoids in the first place. Flanders (1966), in
his excellent discussion of species replacement among the parasitic Hymenoptera, pro-
vided several examples where one introduced parasitoid of a pest displaced another,
but he did not cite any examples of displacement of native parasitoids of pests.
Rosen & DeBach (1979), in their monograph on Aphytis (all species of which are
primary parasitoids of armored scales), predicted that, on average, there should be one
species of Aphytis for each species of armored scale. There are more than 130 species
of armored scales recorded from Florida (Dekle 1976), and hence we should expect a
large number of species of this aphelinid genus particularly from native diaspidids for
which there were no prior records of parasitism by Aphytis. Over the past 7 years,
aided by Dr. David Rosen during his sabbatical in 1991, I collected, and held for
parasitoid emergence, diaspines from many native as well as adventive plants. Although
we reared specimens of this genus from many of the scales, most proved to be introduced









Insect Behavioral Ecology '92: Bennett 55

species of Aphytis or species previously recorded from other scales. Whether or not
introduced species displaced native ones, or whether there were no specialized native
ones attacking many species of the scales, is debatable. The natural enemies of scales
on most plants other than citrus had not been studied before the introduction of Aphytis
holoxanthus DeBach and A. lingnanensis Compere (Hymenoptera: Aphelinidae).
Hence we have no basis to determine whether native species of Aphytis have actually
been displaced. We are left with anecdotal accounts about the partial or complete dis-
placement of native parasitoids by introduced parasitoids. There are examples where a
native parasitoid, which turned its attention to an adventive pest, may have been dis-
placed on that particular host by an introduced parasitoid. There are several examples
wherein one, or more, introduced parasitoids has been displaced by a further introduct-
ion. By reviewing some of these examples perhaps we can deduce whether in fact
native parasitoids are likely to have been completely displaced (eradicated) by intro-
duced ones. I will refer to examples where an introduced parasitoid eliminated its host,
to examples where at least partial displacement of a native parasitoid of a native host
by an introduced parasitoid occurred and to several examples where introduced
parasitoids were displaced by subsequent introductions.

EXTINCTION OF A HOST

During the final quarter of the last century and the first quarter of the current
century, a small moth, Levuana iridescens Bethune-Baker (Lepidoptera: Zygaenidae),
virtually destroyed the coconut industry in Fiji. Although at the time it was not known
to occur elsewhere, and subsequently was not found during extensive surveys for nat-
ural enemies in the Pacific islands and southeast Asia, it was concluded that this moth
was not native to Fiji (Tothill et al. 1930). These authors remark on the rarity of
parasitoids of it in Fiji and use this as one of the criteria in concluding that the moth
was not native. Only three specimens of hymenopterous parasitoids had been reared
from Levuana pupae. Two of the specimens were not adequate for identification beyond
the family level (Chalcididae). The third specimen (also Chalcididae) escaped! Before
escape, it was identified tentatively as a species of Brachymeria known to be parasitic
on the pupa of another moth and therefore likely to be polyphagous on lepidopterous
pupae.
The tachinid Bessa remote (Aldrich) is a parasitoid of the coconut moth Cathartona
catoxena (Hampson) in Malaysia. When introduced into Fiji in 1925, it attacked the
Levuana moth so successfully that the Levuana moth became extinct (Robinson 1975,
Howarth 1991). Hence, had there been specialized monophagous parasitoids of Levuana
moth in Fiji, they would also have become extinct! Perhaps, if indeed the Levuana moth
had originated elsewhere, it has been eliminated in its area of origin by the arrival of
B. remota in the same dramatic manner that occurred following the deliberate introduct-
ion of this tachinid into Fiji. This could account for lack of success in locating the area
of origin of the moth. Heteropan dolensDruce, an unrelated zygaenid moth, disappeared
from Fiji about the same time apparently due to suppression by B. remota (Robinson
1975, Howarth 1991). Fortunately, H. dolens still occurs in Aneityum Island, New
Hebrides (Robinson 1975).
Another example of the elimination of an immigrant host by an introduced polyphag-
ous parasitoid is that of the nigra scale in California. Flanders (1959, 1966) described
the discovery in California of the nigra scale, Saissetia nigra (Nietner), and its disap-
pearance following the introduction of an encyrtid parasitoid, Metaphycus helvolus
Compere, which also attacks several other soft scales.
Howarth (1991), while reviewing the negative impacts of classical biological control,
cited known examples where extinctions of the nontarget as well as the target species









56 Florida Entomologist 76(1) March, 1993

have been reported. These, in addition to the zygaenid moths cited above, include the
sharp decline of native pentatomids in the genera Coleotichus and Oechalia in Hawaii
after the introduction of the tachinid Trichopoda pilipes (F.) and the scelionid Trissolcus
basalis (Wollaston) in 1962 for the control of the immigrant southern green stink bug
Nezara viridula (L.). Also, apparently as a result of these introductions, another immi-
grant pentatomid Murgantia histrionica (Hahn) as well as its deliberately introduced
parasitoid Trissolcus murgantiae (Ashmead) became extinct in Hawaii. Additionally
the disappearance of at least 15 species of the larger native moths of Hawaii was attrib-
uted aby Howarth (1991) to the direct or indirect impact of biological control introduc-
tions, though this is disputed by Funasaki et al. (1988).
DISAPPEARANCE OF A NATIVE PARASITOID

Although the following examples are not conclusive and have not resulted in extinc-
tion, they suggest that competitive suppression of native parasitoids does occur.

Sugarcane Borers

The Caribbean. In the Caribbean and much of Latin America, the most concerted
effort in the field of biological control has been against the sugarcane borer Diatraea
saccharalis (F.) and related species. In most of the Caribbean islands, both pest and
natural enemies are adventive species. However, in Trinidad, four economically impor-
tant species of Diatraea are considered native; several native parasitoids also occur
there. Despite attempts to introduce additional control agents, the only species known
to have become established permanently is the Asian braconid Cotesia flavipes Came-
ron, a parasitoid of species of Chilo, a genus closely allied to Diatraea. Levels of
parasitism of Diatraea spp. in sugarcane by C. flavipes are generally low, and there
appears to be little likelihood that, in Trinidad, it will have much effect on the dominant
native parasitoid Paratheresia claripalpis Wulp. This is not so in maize, where Diatraea
lineolata Wlk., the most abundant stalk borer, is not heavily parasitized by P. claripal-
pis. Until the introduction of C. flavipes, the dominant native parasitoid was Apanteles
diatraeae Muesebeck, a braconid which seldom parasitized Diatraea spp. in sugarcane.
Levels of parasitism of D. lineolata by A. diatraeae reported in the earlier literature
(Kevan 1945) seldom exceeded 10%, although levels of 30% were found in later surveys
(Bennett unpubl.). However, after the establishment and build-up of C. flavipes in
Trinidad, parasitism by A. diatraeae diminished to the point where it was not rep-
resented in extensive collections of D. lineolata during 1984-85 (Bennett unpubl.).
Brazil. In Brazil occur species of Diatraea additional to D. saccharalis. An additional
native tachinid, the Amazon fly, Metagonistylum minense Townsend, is widespread in
Brazil. In 1974, C. flavipes was introduced into northeast Brazil where it established
readily on Diatraea flavipennella Box and D. saccharalis; however, when it was re-
leased in Sa6 Paulo State, it barely established, and required frequent supplementary
releases to have any effect on borer populations. A cool-weather strain was acquired
from Pakistan in 1978; it performed well and is now the dominant parasitoid (Botelho
1992). The native tachinid parasitoids Metagonistylum minense and P. claripalpis have
become scarce. While they are no longer represented in survey collections in many
fields they occur sporadically in collections from other fields.

Florida Red Scale

Florida. The success story of the biological control of the Florida red scale Chrysom-
phalus aonidum L. has been chronicled several times (Selhime et al. 1969, Browning
1990). Considered to be one of the most serious pests of citrus, Florida red scale was









Insect Behavioral Ecology '92: Bennett


brought under excellent biological control by the introduction of Aphytis holoxanthus
DeBach (Aphelinidae) in 1960. By 1964, scale populations had been reduced to a non-
economic level. Selhime et al. (1969) report that it successfully replaced
Pseudhomalopoda prima Girault (Encyrtidae) as the main control agent. Displacement
was so complete that it is often difficult to find this encyrtid in Florida red scale on
citrus. Despite its common epithet, the Florida red scale is not native to Florida,
whereas P. prima is. P. prima is restricted to the neotropics and to the southern States
from Florida west to Texas and the host range is given as C. aonidum and Aonidiella
aurantii Maskell, both citrus pests of Asian origin. Therefore, even if P. prima disap-
peared completely from the citrus ecosystem (which is not a native ecosystem in Florida,
and P. prima hasn't disappeared completely) it would merely be a retreat to its native
host or hosts. I have reared it as the dominant parasitoid of Acutaspis morrisonorum
Kosztarab on southern red cedar, Juniperus silicicola (Small) Bailey, as recently as
1990. Hence, although it has been displaced to a great extent by A. holoxanthus on
Florida red scale, it still is the most important parasitoid on at least one of its native
hosts.
Brazil. Following its success in the USA, A. holoxanthus was colonized, in 1962 by
Paul DeBach, in the State of Sa6 Paulo, where previously the native aphelinid Aphytis
costalimai (Gomes) was the most common parasitoid (Rosen & DeBach 1979). In 1984,
while in Brazil to collect A. costalimai for dispatch to India for trial on Melanaspis
glomerata (Green), I examined several thousand Florida red scale; several hundred
Aphytis pupae were obtained. The following excerpt of my unpublished tour report
explains the findings:
"A few of the parasites emerging from Chrysomphalus aonidum were slide-mounted
in Hoyer's. None proved to be A. costalimai, a species readily recognizable by its
distinctively mottled fore-wing. Similarly, none of the pupae examined appeared to be
as extensively pigmented as those of A. costalimai (Rosen & DeBach 1979). The speci-
mens proved to be Aphytis holoxanthus. The ease with which A. costalimai was encount-
ered in the past on this host (Rosen & DeBach 1979) suggests that, if present, it should
have been represented in the sample size obtained. If the Aphytis material shipped to
India proves to be A. holoxanthus, a species introduced into Brazil for the control of
Florida red scale (Rosen & DeBach 1976), it is evident that this species has completely
displaced A. costalimai as a parasite of C. aonidum. In view of the relative scarcity of
C. aonidum, A. holoxanthus appears to have effected excellent control." (F. D. Bennett
1984 unpubl. report ).
All specimens of Aphytis shipped to India were indeed A. holoxanthus. In 1987 I
reared a few specimens of A. costalimai from Lindingaspis sp. (Diaspididae), collected
on Ligustrum sp. at Curitiba, Parana, Brazil proving conclusively that this parasitoid
had not been driven to extinction.

DISPLACEMENT OF INTRODUCED PARASITOIDS BY OTHERS

Rhodesgrass Mealybug

The Rhodesgrass mealybug, Antonina graminis (Maskell), a pest of Asian origin,
threatened to ruin the cattle industry in Texas in 1945 because of its deleterious effect
on pasture grasses. Excellent biological control was achieved following the introduction
of parasitoids (Dean et al. 1979). The mealybug, following its discovery in 1945, also
was considered a serious pest in Florida, where it attacked a wide range of grasses
(Questel & Genung 1957). Biological control efforts in Florida commenced with the
introduction of Anagyrus antoninae Timberlake (Hymenoptera: Encyrtidae) in 1954
(Questel & Genung 1957). In July, parasitoids obtained from Texas were released in the
Clewiston area, and in November in the Homestead area (Questel & Genung 1957, Dean









58 Florida Entomologist 76(1) March, 1993

& Schuster 1958). In 1956, parasitoids were recovered several miles from the nearest
release sites. Some were distributed to new areas by collecting grass, infested with
parasitized mealybugs, and placing it in areas where parasitoids did not occur. In areas
where parasitoids were released first, the mealybug had become scarce by 1957. Questel
& Genung (1957) noted that establishment had occurred at every release site, and
considered that the continued dissemination of this parasitoid would aid in controlling
Rhodegrass mealybug throughout southern Florida.
The introduction into Florida of Neodusmetia sangwani (Subbo Rao), the encyrtid
parasidoid which provided successful control of Rhodesgrass mealybugs in Texas, were
recommended. Large-scale releases of this parasitoid were made in 1959. Although
establishment occurred, there are no published accounts of a thorough post-release
survey. The late Professor Reece I. Sailer collected extensively and reared parasitoids
of Rhodesgrass mealybug from 1975-1985. During 1985-1991 I continued to collect and
rear parasitoids from localities throughout Florida. N. sangwani was widespread and
abundant. In contrast, A. antoninae was never found during Sailer's or my surveys.
These results suggest strongly that A. antoninae, which initially established readily
and showed promise as an effective control agent, has been displaced completely by N.
sangwani. Another encyrtid Pseudectroma sp. of unknown origin (not P. europaea
(Mercet) which had been introduced as a control agent), was encountered frequently
and, although never as abundant as N. sangwani, might also have been a contributory
factor to the disappearance of A. antoninae. Schuster & Dean (1976) reported the
competitive displacement of A. antoninae by N. sangwani in Texas and suggested that
the "lack of competitiveness of A. antoninae was a result of its inability to develop at
high vapor deficits" and at high seasonal temperatures prevailing during the summer
months. They found that A. antoninae was displaced but not eliminated. In Florida
despite a more equitable rainfall distribution A. antoninae appears to have been driven
to extinction or at least to a non-detectable level.

Fruitflies in Hawaii.

The Oriental fruitfly (Bactocera (= Dacus) dorsalis (Hendel)) and the Mediterra-
nean fruitfly (Ceratitis capitata (Wiedemann)) both became serious pests after their
arrival in Hawaii. Three of several braconid parasitoids introduced in rapid succession
in 1948 for the control of the Oriental fruit fly became established. Initially Biosteres
(= Opius) longicaudatus (Ashmead) became widely established, but eventually Bio-
steres vandenboschi and finally Biosteres arisanus (Sonan) (= Opius oophilus Fullaway)
became dominant to the extent that the other two species have been relegated to the
status of rare species in some of the islands (Van den Bosch et al. 1982). B. arisanus
now accounts for 74 to 92% of all parasitoids reared from B. dorsalis and C. capitata
in both tropical and temperate zones in Hawaii (Ramadan et al. 1992). This provides us
with an example of competitive displacement of introduced parasitoids of immigrant
pests by another introduced parasitoid. However, after forty years, the other
parasitoids are still well represented in samples and there is no apparent likelihood that
any species will be driven to extinction.

Competition Among Aphytis Species

Perhaps the most conclusive evidence for competitive displacement and likelihood
of extinction over large areas is to be found in the extensive studies on Aphytis spp.
by DeBach & Sundby (1963). Aphytis chrysomphali (Mercet) was eliminated from
nearly all of its range (4,000 sq. miles) in southern California within 10 years of the
introduction of A. lingnanensis which in turn was displaced over much of its range









Insect Behavioral Ecology '92: Bennett


within 4 years by A. melinus DeBach from India. However, A. lingnanensis precluded
the establishment of A. melinus in the milder climatic areas of San Diego County. In
some areas, host scarcity was not a limiting factor. In laboratory studies, whenever any
two of three species (A. fisher DeBach, A. melinus, and A. lingnanensis) were cul-
tured together, one species was eventually eliminated: the species surviving was influ-
enced by the temperature and humidity regimes of the particular experiment. Even
when an abundance of hosts was present at all times, A. lingnanensis eliminated A.
fisher after 9 generations. While the laboratory studies largely confirmed field events,
A. chrysomphali was able to compete successfully and to coexist with A. lingnanensis
in a few coastal areas where conditions for its survival were optimal.
The elimination of one parasitoid by another under laboratory conditions has been
demonstrated. Using her ovipositor, the female of A. lingnanensis mutilated the pre-
pupae and pupae of Encarsia lounsburyi (Berlese & Paoli) (Flanders 1951). Similarly,
in laboratory culture with black scale as host, Coccophagus pulvinariae Compere
(Aphelinidae) displaced Coccophagus cowperei Girault. However, in the field both were
able to co-exist because the host preference of the females was not the same (Compere
1940). DeBach (1965, 1966) discussed competitive displacement of ecological homologues;
included among the factors accounting for displacement were temperature, humidity,
natural enemies, disease, type or condition of the food source as well as other factors.
He dealt with numerous examples including those where one parasitoid succeeded
another and stated, in the case of the Aphytis spp. introduced for control of California
red scale referred to earlier, that displacement could occur in the presence of "surplus,
even abundant, food". Huffaker & Laing (1972) attempted to clarify this statement and
concluded that "competitive displacement must involve competition for some requisite
(a resource or relatively enemy-free space), however low its intensity".

INEFFECTIVE INTRODUCED PARASITOIDS, EFFECTIVE
IMMIGRANTS, AND REFUGIA

Introduced parasitoids do not always dominate native species (DeBach 1966). In
fact, in classical biological control, failure to obtain establishment at all is frequent (Hoy
1985). DeBach (1965) stated that perhaps only 25% of imported colonized entomophag-
ous insects become established. In other instances establishment occurs but there is
only minor effect on the host or on native parasitoids. The introduction of parasitoids
and predators has reduced host densities adequately to replace chemical pesticides in
approximately 16% of more than 600 projects (Myers et al. 1989). The reasons for
non-establishment of introduced parasitoids, in most instances, remain unclear and have
been debated repeatedly by biological control practitioners, ecologists and modelers
(e.g., Hoy 1976, Ehler & Hall 1982, 1984, Keller 1984, Myers et al. 1989 and DeBach
& Rose 1992). Competitive exclusion is a frequently debated issue, but in their analysis
Elher & Hall (1982, 1984) and Keller (1984) were considering exotic insect introductions
and not native natural enemies.
Tallamy (1983), when analyzing the results of the numerous parasitoid introductions
into the USA for the control of the gypsy moth, Lymantria dispar (L), in the context
of equilibrium biogeography notes that "parasite extinction events are regular occur-
rences".
Assessment prior to the intensive programs for the importation of parasitoids from
Europe indicated that the level of parasitism of gypsy moth by native species was
negligible (Howard & Fiske 1911, Burgess 1926). If these native species were not de-
tected in post-introduction surveys for gypsy moth parasitoids they should not necessar-
ily be judged as having been driven to extinction. In all probability they retreated to
the hosts which they attacked before the arrival of the gypsy moth. It is possible that









Florida Entomologist 76(1)


certain of the introduced parasitoids could have had a negative impact on native
parasitoids on their native hosts. For example Brachymeria intermedia (Nees)
(Hymenoptera: Chalcididae) is polyphagous in its native range, where it parasitizes at
least 15 species of Lepidoptera. Following its introduction into the USA it has been
reared from several leaf rollers as well as from gypsy moth (Leonard 1981). Similarly
the introduced tachinid fly Compsilura concinnata (Meigen) has been reared from sev-
eral hosts including non-forest dwelling species such as the imported cabbage worm
Pieris rapae (L.) and the cabbage looper Trichoplusia ni (Htibner) in eastern USA
(Headland & Schroeder 1981). The three species of native Tachinidae reared from gypsy
moth (Howard & Fiske 1911) all have a very broad host spectrum (see Arnaud (1978)
for current nomenclature and recorded hosts). Hence introduced parasitoids are unlikely
to displace them over their entire host range.
There are several other examples in the literature where successive introductions
of additional parasitoids have led to improved biological control. In some instances, the
parasitoid introduced earlier became scarce and the more recently introduced species
provided effective control. In others, the second introduction complemented the first
and both species co-existed, as occurred during the olive scale biological control program
in California (Huffaker & Kennett 1966). In several biological control programs, the
successive introduction of different biotypes has led to improved control (Caltagirone
1985, LaSalle & Gauld 1992) and to the apparent disappearance of the first biotype.
This may have occurred more frequently than reported, but was not recognized because
of the lack of criteria to differentiate between successive introductions of the same
species carrying a different genome.
Introduced parasitoids do not necessarily compete well with native species. In ex-
perimental releases of parasitoids in feed lots in northern Nebraska for fly control, the
native pteromalid Muscidifurax raptor Kogan & Legner outperformed the introduced
Pachycrepoideus vindemiae (Rondani) (Hymenoptera: Pteromalidae) (Petersen et al.
1992).
Examples heretofore given of adventive parasitoids have dealt with introduced
species. There is necessarily much less documentation of immigrant species, i.e., those
which arrived as stowaways or hitchhikers or even by flight (Simberloff 1986). This is
because there is no record of immigrant parasitoids which fail to establish.
Parasitoids sometimes do immigrate. In Hawaii, studies by Sembel (1980) suggested
that the immigrant Trichogramma papilionis Nagarkatti, first recorded on Oahu in
1979, multiplied rapidly. It became dominant over Trichogramma chilonis Ishii as an
egg parasitoid of several but not all lepidopterous hosts. Hirose et al. (1988), when
discussing their studies on the coexistence and interspecific competition of three species
of Trichogramma as parasitoids of the swallowtail Papilio xuthus L. in Japan, con-
cluded that habitat differences of the parasitoids over the entire range of the hosts
precluded any one of the three species being driven to extinction.
In short, in addressing the question of whether introduced parasitoids replace native
parasitoids, there is, as is true for most other biological phenomena, no unequivocal
simple answer. The examples cited above suggest that sometimes native parasitoids
can be displaced over much of their range, but usually there are restricted habitats or
refugia where they are able to co-exist with introduced parasitoids. Where host extinc-
tions do occur they are more likely the result of the introduction of a generalist
parasitoid like Bessa remota that can survive on alternate hosts than from the introduc-
tion of host-specific parasitoids.

ACKNOWLEDGMENTS

I wish to thank J. H. Frank for inviting me to participate in this Symposium and to
acknowledge especially his assistance and suggestions in completing the manuscript. M.


March, 1993









Insect Behavioral Ecology '92: Bennett


A. Hoy and E. D. McCoy also reviewed earlier drafts of the manuscript and offered
valuable suggestions for its improvement. This is University of Florida, Institute of
Agriculture and Food Sciences Journal Series No. R-02708.

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Florida Entomologist 76(1)


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March, 1993









Insect Behavioral Ecology '92: Bennett 63

antoninae (Hym.: Encyrtidae) by its ecological homologue Neodusmetia
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VAN DEN BOSCH, R., P. S. MESSENGER, AND A. P. GUTIERREZ. 1982. An introduc-
tion to biological control. Plenum Press, New York. 247 p.












ANT BEHAVIOR AND MICROBIAL PATHOGENS
(HYMENOPTERA: FORMICIDAE)

DAVID H. OI' AND ROBERTO M. PEREIRA
Entomology & Nematology Department
University of Florida
P.O. Box 110620
Gainesville, FL 32611-0620

'Current Address: USDA-ARS Medical & Veterinary Entomology Research Lab.,
P.O. Box 14565,
Gainesville, FL 32604

ABSTRACT

The effectiveness of microbial controls for pest ants can be reduced by ant behaviors.
Introductions of pathogens, including nematodes, into ant nests result in behavioral
responses by ants that affect infection rates to ants exposed to inocula, affect the dis-
semination of inocula among nestmates, and affect the dispersal of inocula outside the
nest. These behaviors include grooming, secretion of antibiotics, nest hygiene, avoid-
ance, and dispersal. Ant behaviors must be considered in developing microbial control
agents. Approaches to overcoming the behavioral responses of the red imported fire
ant to the entomopathogen Beauveria bassiana (Balsamo) Vuillemin are discussed.









64 Florida Entomologist 76(1) March, 1993

RESUME

La eficiencia del control microbiano contra hormigas plaga puede ser reduzida devido
al comportamiento de las hormigas. La introducci6n de pat6genos, incluyendo neAm-
todos, en el nido de las hormigas Ileva a cambios comportamentales que afectan la
porcentaje de infecci6n de las hormigas expuestas al in6culo, la dispersi6n del in6culo
entire las hormigas de un mismo nido, y la dispersi6n del in6culo fuera del nido. Estos
comportamientos incluyen limpieza del cuerpo de las hormigas, secreci6n de antibi6ticos,
higiene del nido, el evitar al pat6geno, y dispersion de las hormigas. El comportamiento
de las hormigas deve ser considerado durante el desarrollo de agents microbianos para
control biol6gico. Propuestas para superar las respuestas comportamentales de la "hor-
miga brava" al entomopat6geno Beauveria bassiana (Balsamo) Vuillemin.





Ants, as social insects, have many unique behavioral adaptations that facilitate their
survival. Many of these adaptations have frustrated man's attempts to control ants by
various means. Understanding these adaptations is especially important when the use
of biological control agents is desired. The successful control of pest ants through the
use of parasitoids, predators, or pathogens has not been documented, although natural
enemies do exist (Wojcik 1989, Jouvenaz 1983). Perhaps the lack of success or even
attempts at biological control may be partly attributable to the behavior of ants. Their
ability to fend off parasitoids to protect scale insects (DeBach 1974), or their ability to
defend themselves against phorid flies (Wojcik 1989), are indicative of the inherent
difficulty in finding effective biological control agents for ants. Behavioral responses to
the presence of natural enemies may be especially important when pathogens are used
as microbial insecticides, in contrast to the use of classical biological control methods.
Behavioral responses and interactions of ants with their arthropod parasites were re-
viewed by Wojcik (1989). With respect to microbial pathogens, Wheeler (1910) ques-
tioned whether ants could be controlled with pathogens given the environment that
they inhabit. He reasoned that ants had evolved mechanisms that prevented their nests
and themselves from being overrun by fungi and other microorganisms. However,
Evans (1982) disputed reports (Allen & Buren 1974) that fungal epizootics of ants are
uncommon, and has reported extensively on Cordyceps and other fungi parasitizing ants
in the tropics. Thus, the allure of finding a microbial biological control for pest ants is
still present as evidenced by recent research in this area by Drees et al. (1992), Jouvenaz
& Martin (1992), Pereira & Stimac (1992), Pereira et al. (1993a, 1993b), SAnchez-Pefia
& Thorvilson (1992), Siebeneicher et al. (1992), and Stimac et al. (1993) who examine
the potential of controlling the red imported fire ant, Solenopsis invicta Buren, which
immigrated into the southeastern U.S. over fifty years ago. Yet, these recent works
again confirm the difficulty of obtaining control with this strategy.
In this paper, we review examples of reported ant behaviors that occur in the
presence of pathogens, including nematodes (Table 1). While these behaviors may not
have evolved specifically in response to microbial antagonists, they have negative and
positive effects on the efficiency of infection and spread of disease. We have categorized
these behaviors as affecting: 1) infection of ants exposed to inoculum; 2) intracolony
dissemination of inoculum among nestmates; and, 3) dispersal of inoculum outside the
nest (Table 2). The implications of these behaviors on the development of microbial
control agents for imported fire ants will then be discussed, using our research on
Beauveria bassiana as an example.










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Florida Entomologist 76(1)


TABLE 2. SOME EXAMPLES OF ANT BEHAVIORS AND THEIR EFFECT IN INCREASING
(+) OR DECREASING (-) INFECTION RATE, DISSEMINATION OF THE
PATHOGENS WITHIN THE ANT NESTS OR DISPERSAL OF PATHOGEN OUT-
SIDE OF THE NESTS.

Intra-Colony Out of Nest
Behavior Infection Dissemination Dispersal

Grooming + ne2
Use of Secretions ne
Avoidance of Pathogens ne
Nest Hygiene/ Necrophoresis ne -/+ -/+
Summit Disease Syndrome ne +
Altered Activity Time or Place ne -/+
Colony Movement ne +

'This effect reported for termites (Kramm et al. 1982), but not for ants.
'ne = no effect



GROOMING

Self-grooming and mutual grooming by any nestmates can either impede or facilitate
the infection and spread of disease. Wilson (1971) described the grooming process as
involving the use of tibial combs, the rubbing of legs, and licking to keep body surfaces
clean. Inocula are not ingested because they are collected in an infrabuccal pocket, a
cavity on the ventral surface of the buccal chamber (Eisner & Happ 1962), and expelled
as a pellet with other debris (Siebeneicher et al. 1992). In the fungivorous leaf cutting
ant Acromyrmex octospinosus, labial gland extracts containing chitinases were
hypothesized to have a fungistatic function against microflora in the infrabuccal pocket
(Febvay et al. 1984).
When grooming removes inoculum from the cuticle before infection occurs, it obvi-
ously reduces infection rates. Observations at 2 h intervals with scanning electron micro-
scopy of S. invicta inoculated with B. bassiana conidia has demonstrated the removal
of conidia from the integument of adults and larvae (unpublished data) (Fig. 1).
Siebeneicher et al. (1992) reported similar observations, and SAnchez-Pefa & Thorvilson
(1992) inferred that grooming prevented infection from Conidiobolus conidia. Likewise
another myrmicine ant, Cephalotes atratus removed spores of a Cordyceps sp. by
grooming (Evans & Samson 1982). Drees et al. (1992) reported the incessant preening
by red imported fire ant workers of brood, alates, and themselves after exposure to
Steinernema and Heterorhabditis nematodes, in an apparent attempt to remove them.
While grooming may reduce infection by the removal of inocula, grooming might
assist in the dissemination of inocula among nestmates. The transfer of inocula to work-
ers may occur by incidental contact during the grooming of contaminated nestmates.
While this has not been reported for ants, dissemination of inocula during grooming has
been seen in termites (Kramm et al. 1982).

USE OF SECRETIONS

The use of antimicrobial secretions is another behavior that reduces the chance of
infection once inoculum is present on the ant. Venom from S. invicta includes antibiotic
alkaloids (Blum et al. 1958, Jouvenaz et al. 1972, Storey 1990) that are sprayed during
gaster flagging behavior (Obin & Vander Meer 1985). This behavior included the raising


March, 1993












Insect Behavioral Ecology '92: Oi & Pereira 69













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Florida Entomologist 76(1)


and wagging of the gaster to provide a directed spray onto brood, and also directed
against other ants during heterospecific confrontations. Storey (1990) demonstrated the
fungistatic activity of venom alkaloids and recorded increased levels of these products
in nest soil containing B. bassiana conidia. This suggested that venom release could be
induced by the presence of this entomopathogen.
Besides the venom alkaloids, secretions from the metapleural glands of Myrmica
laevinodis Nyl., Myrmica rubida (Latreille), Atta sexdens, Acromyrmex sp., Messor
barbarus L., and Myrmecia nigriscapa were shown to have antibiotic effects against
bacteria and fungi (Maschwitz et al. 1970, Schildknecht & Koob 1971, Beattie et al. 1985,
1986). In the case of the Myrmica species, these secretions, which contain phenyl acetic
acid and/or myrmicacin (1-p-hydroxydecanoic acid) (Schildknecht & Koob 1971), are
spread over the cuticle by self grooming and nestmate grooming. Secretions also were
spread over larvae (Maschwitz et al. 1970) presumably to protect these individuals from
diseases. However, Beattie et al. (1986) observed the flowing of secretions along the
grooves of an ant's integument, perhaps indicating that grooming was not always neces-
sary to distribute the secretions.


NEST HYGIENE

Nest conditions, such as high humidity and relatively stable temperatures, favor
growth of microbial pathogens. Thus, behaviors that limit the occurrence of epizootics
within a colony are vital for survival. The elimination of potential sources of inocula is
an obvious benefit of nest hygiene. This occurs regardless of the presence of pathogens
and is evidenced by kitchen middens in the vicinity of nests. Necrophoresis, or cadaver
removal, has been seen in S. invicta infected with B. bassiana (Storey 1990,
Siebeneicher et al. 1992, Pereira & Stimac 1992), and in C. atratus, infected with
Cordyceps sp. (Evans & Samson 1982, Evans 1982). We have observed piles of spor-
ultaing B. bassiana infected cadavers of S. invicta outside of nests. These sporulating
piles promoted dispersal of the pathogen. However, this external nest dispersal appar-
ently does not occur in the case of Formica rufa adults infected with an unknown
pathogen, where their removal and supposed consumption by healthy nestmates was
reported (Marikovsky 1962). Consumption of infected individuals, however does not
always preclude the dispersal of pathogens. In Solenopsis geminata, spores of the
protozoan Burenella dimorpha are transferred to fourth instar larvae when they are
fed infected pupae cannibalized by workers (Jouvenaz 1983).
Another aspect of nest hygiene would include the isolation of infected cadavers
within the nest. S. invicta workers pack soil around dead individuals infected with B.
bassiana (Storey 1990), and this behavior contributed to the lack of transmission of
fungal disease within artificial fire ant nests in the presence of soil (Pereira & Stimac
1992). From these results, one may infer that the isolation of infected cadavers with
nest soil may provide an effective barrier to the spread of B. bassiana.


AVOIDANCE OF PATHOGENS

In addition to the removal or isolation of inoculum sources, the simple avoidance of
inocula is another behavior that limits the spread of infections. Marikovsky (1962) re-
ported the avoidance by F. rufa of infected nestmates covered with conidia. These
cadavers were contagious, and were not removed or consumed, in contrast to
nonsporulating, infected individuals. Similarly, during bait testing, Zoebisch & Stimac
(1990) reported that S. invicta avoids baits containing B. bassiana conidia.


March, 1993









Insect Behavioral Ecology '92: Oi & Pereira


COLONY MOVEMENT

Field applications of nematodes to S. invicta nests resulted in the movement of at
least a third of the treated colonies to new nest sites (Drees et al. 1992). We observed
a similar tendency of nest relocation for fire ant colonies injected with conidia of B.
bassiana (unpublished data). These colony movements probably reflect an avoidance by
the ants from further contact with the nematodes or Beauveria conidia.

DISPERSAL

The most frequently recorded behavioral response to microbial pathogens is the
dispersal of ants infected with fungal pathogens (Table 1). Whether the prevalence of
this behavior in the literature is a reflection of its actual frequency of occurrence in ant
populations, or whether it is just a function of its conspicuousness, is not clear. In
general, infected ants were reported to disperse to the tops of grass blades or other
vegetation, and then die clinging to the vegetation with their legs and mandibles. Some
fungi, fasten the ants firmly to the substrate by fungal outgrowths, or rhizoids (Evans
1974, Balazy & Sokolowski 1977, Humber 1981). Subsequently fruiting bodies will
emerge from the cadaver, whose elevated location would facilitate dispersal of inoculum.
Loos-Frank & Zimmerman (1976) attributed the propensity to climb in ants infected
with Pandora (Erynia = Entomophthora) formicae, to the growth of hyphae within the
nervous system.
In addition to dispersal to elevated locations, ants with fungal infections have been
observed to leave their nest while displaying erratic, spasmatic movements (Marikovsky
1962, Evans 1982). In arboreal ant species, dispersal was toward the ground, where
individuals were found under leaf litter (Evans 1982). Uncharacteristic behavior also
has been reported in Camponotus castaneus infected with the nematode Rabbium
paradoxus; infected individuals of this nocturnal ant became diurnal (Poinar et al. 1989).
In laboratory experiments with small S. invicta colonies, 3 to 5 days after application
of B. bassiana, infected ants were observed wandering outside of the nest cells.
Likewise, in field injections of B. bassiana into fire ant nests, we observed erratic
movements of masses of ants around the nests also within 3 to 5 days of exposure
(unpublished data). Contrary to their typical behavior of walking on the surface of the
ground, these undisturbed fire ants moved to the tops of the blades of grass and appar-
ently were not foraging or alarmed. Such behavior may be related to the fever responses
or "summit disease syndrome", reported for fungus-infected grasshoppers and caterpil-
lars which climb up vegetation to sun themselves to increase body temperatures and
eliminate the fungal infection (Marikovsky 1962). However, despite the many examples
of upward dispersal of infected ants, the elimination of infection has not been reported.
Regarding the dispersal of infected ants away from the nest, some have speculated
on the altruistic nature of this behavior, since potential sources of inocula are removed
from the nest (Evans 1982, 1989). If infected ants die within their nests, either they
would be removed from the nest, or sequestered within the nest before the pathogen
reached the infective stage. Thus, dispersal behavior does not seem to be a major factor
in mitigating colony exposure to the pathogen, unless infection rates are so rapid and
extensive that the hygienic response of the ants is overwhelmed. However, ant dispersal
may aid in inoculum dissemination outside a nest, and thus is a benefit to the pathogen,
especially if the pathogen is not an obligate parasite of a single host.
Table 2 summarizes the positive or negative effects of the ant behaviors discussed
above on infection rate and dissemination of pathogens. Published reports on the effec-
tiveness of different ant behaviors inhibiting the development of diseases tend to be
anecdotal. Studies quantifying the effects of ant behavior on rates of infection under









Florida Entomologist 76(1)


field conditions would be difficult to conduct. Nevertheless, the value of these behaviors
in disease prevention is attested to by the fact that (1) a majority of the behavioral
responses of ants to pathogens are described as limiting infection or pathogen dissemi-
nation, and (2) relatively few examples of disease epizootics are reported for ant popu-
lations.

IMPLICATIONS OF ANT BEHAVIORAL RESPONSES TO MICROBIAL PATHOGENS
ON THE DEVELOPMENT OF MICROBIAL CONTROL AGENTS FOR ANTS

The responses of ants to microbial pathogens must be considered in developing
microbial control agents for ants. Research with B. bassiana for imported fire ant
control can be used as an example of some of the implications of the behavioral responses
of ants toward the development of a microbial control.
Stimac et al. (1989) demonstrated the pathogenicity of an isolate of B. bassiana
recovered from fire ants in Brazil to Solenopsis spp. This isolate was brought to Florida
for further study as a potential biological control agent for imported fire ants in the
U.S. Initially, the idea was to use the fungus as an inoculative introduction that would
initiate naturally occurring epizootics in ant nests. However, after considering the re-
sults of initial studies, and the effects of ant behaviors on the performance of the fungus
(Pereira & Stimac 1992, Pereira et al. 1993a, 1993b), it became clear that two major
problems related to ant behavior needed to be addressed: 1) overcoming the behaviors,
such as grooming, that prevent infection when individual ants are exposed to inoculum;
and, 2) bypassing the negative effects of behaviors, such as necrophoresis and dispersal
of infected ants, that prevent transmission among nestmates.
A potential approach toward overcoming the first problem is to improve contact
between the ant cuticle and conidia by finding a formulation that is resistant to groom-
ing. As an added measure, a large developing dose of conidia needs to be delivered to
the ants to allow some of the conidia to escape removal by grooming, and inactivation
by antibiotic secretions. To overcome the lack of nestmate transmission, a mass inocu-
lation was sought by injecting conidia directly into the nest. Provided that the injection
application obtained good dispersion throughout the nest, the need for transmission
among nestmates could be eliminated. Given good dispersal throughout a nest, and a
formulation that cannot easily be groomed off by the ants, it should be possible to obtain
a mass infection that would overwhelm the hygienic response of ants, disrupt social
order, and eliminate the colony.
In conclusion, ant behaviors affect both directly and indirectly the performance of
pathogens within the nests. These behaviors may or may not have evolved in response
to challenges by pathogens or due to interactions of the pathogens with the neurological
system of the ants. Although some behaviors favor the dispersal of inocula, most of the
reported behaviors are detrimental to the pathogens and may frustrate us in our at-
tempts to develop microbial controls for pest ants. The ants' amazing and fascinating
abilities to adapt and survive have to be understood better if we are ever going to
control pest ants with microbial pathogens.

ACKNOWLEDGMENTS

We thank Mark Deyrup and Earl McCoy for their critical reviews of the manuscript,
and J. Howard Frank and Jon Allen for encouraging our participation in the symposium.
We also appreciated the support of Jerry L. Stimac throughout the course of this work.
This is Florida Agricultural Experiment Station Journal Series No. R-02854.


March, 1993











Insect Behavioral Ecology '92: Oi & Pereira


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(p"









Insect Behavioral Ecology '92 Parkman & Frank


USE OF A SOUND TRAP TO INOCULATE STEINERNEMA
SCAPTERISCI (RHABDITIDA: STEINERNEMATIDAE)
INTO PEST MOLE CRICKET POPULATIONS
(ORTHOPTERA: GRYLLOTALPIDAE)

J. P. PARKMAN AND J. H. FRANK
Entomology & Nematology Department, University of Florida,
P.O. Box 110620, Gainesville, Florida 32611-0620

ABSTRACT

We report on a novel method to disseminate an entomopathogenic nematode,
Steinernema scapterisci Nguyen & Smart, planned to capitalize on the potential of the
nematode as a classical, inoculative biological control agent. Scapteriscus mole crickets
were attracted to synthetic mole cricket song and thus exposed to up to 1.5 million
infective-stage S. scapterisci during the spring and early summer mole cricket flight
season for 2 consecutive years. Releases were made on 28 Florida golf courses and one
sod farm. The nematode was established by exposing attracted mole cricket adults to
infective-stage juvenile S. scapterisci held in moist sand (seven sites) and in damp
upholstery foam (2 sites). Based on capture of nematode-infected mole crickets at least
three months after release, S. scapterisci was established on eight of the golf courses
and the sod farm.

RESUME
Se report un nuevo metodo para diseminar un nematodo entomopatogeno,
Steinernema scapterisci Nguyen & Smart, y se planea hacer emphasis en el potential
de el nematodo como un metodo de control biologico clasico e inoculativo. Durante 2
anos consecutivos en el period de vuelo primaveral y de verano de los grills topos
Scapteriscus, se atrayeron estos a el sonido artificial del grillo y se expusieron a 1.5mil-
lones de estados infectivos de S. scapterisci. Las liberaciones de los nematodos fueron
hechas en 28 campos de golf en Florida y en una finca de 24 cesped. El nematodo S.
scapterisci se establecio al exponer los grills atraidos a los estados juveniles infectivos
del nematodo mantenido en arena humeda (6 lugares) y el espuma sintetica humeda (2
lugares). Basados en la capture de grills topos infectados con el nematodo al menos 3
meses despues de la liberation, se consider que S. scapterisci se establecio en siete
campos de golf y en una finca de cesped.


Scapteriscus mole crickets, immigrant pests of turf and pasture grasses and of veg-
etable seedlings in the southeastern USA, are targets for classical biological control
(Frank 1990). One of the biological control agents under evaluation is the en-
tomopathogenic nematode Steinernema scapterisci (Nguyen & Smart 1990, 1991) from
South America, homeland of Scapteriscus.
Imported into Florida in 1985, S. scapterisci was released experimentally in pastures
in Alachua County, Florida, that same year (Hudson et al. 1988). A single release of
nematodes at each site established a population which persisted until mid- 1990, at
which time monitoring was discontinued (Parkman et al. in press). The persistence of
the nematode at each site and its spread to additional localities (Parkman et al. in press,
Parkman & Frank 1992) showed that it has the attributes of a classical, inoculative
biological control agent. A typical biopesticide, in contrast, would have been expected
neither to persist nor to spread.









Florida Entomologist 76(1)


Largely because its capabilities were unknown in 1985, the nematode was applied
at 200,000 per m2 (2 billion per ha), far exceeding the levels typical for insect parasitoids
when they are used as classical biological control agents, but typical of the use of
entomopathogenic nematodes used as biopesticides. Such high dosage, in retrospect,
undoubtedly was unnecessary to establish populations of the nematode. Far lower num-
bers per unit area might have sufficed to establish populations, but at that time there
had been no reported inoculative use of an entomopathogenic nematode (Kaya 1990).
Adult mole crickets, especially females, are attracted by the song of males, and are
attracted strongly by high-volume synthetic song (Walker 1982). A unique feature of
the 1985 releases was use of synthetic song in attempt to assemble additional mole
crickets to two of the sites inoculated with S. scapterisci (Hudson et al. 1988). After
the success of these first releases, trial releases of S. scapterisci on golf courses were
planned and the concept of using synthetic song was adopted and elaborated upon. The
experimental design of these releases not only takes advantage of being able to attract
mole crickets but also capitalizes on the attributes of S. scapterisci as an inoculative
biological control agent.

MATERIALS AND METHODS

The basic concept of the experimental design was to assemble adult mole crickets
to a concentration of S. scapterisci instead of broadcasting S. scapterisci over large
areas. Thus, 3 million nematodes annually were used per release site instead of 2 billion
per ha in treated areas of each golf course. An emitter producing synthetic sound of the
song of a mole cricket male was used to attract mole crickets to the nematodes. The
basic design of each emitter was an elaboration of the design reported by Walker (1982).
Each emitter contained a sound-synthesizer, an amplifier, a speaker, and a photocell,
with a 12vDC rechargeable power source or a 115v AC/12v DC invertor and a 115v AC
power outlet. The sound-synthesizer was a computer chip programmed with the song
of S. vicinus Scudder and S. borellii Giglio-Tos, either song selectable by an internal
switch within the emitter. Emitters were designed and built by B. J. Mans (Mountain
View, California). In operation, the song selected would be emitted automatically at
105 dB, beginning nightly at sunset and ending two h later. Each golf course was
supplied with two such emitters (to be operated within 2 m of each other), one to play
the song of S. borellii, and the other to play the song of S. vicinus. These emitters
attracted adult mole crickets of the two species during their flight seasons, primarily
in spring but secondarily in autumn (Walker 1985).
Two emitters were set up as part of an infection station to attract flying, adult mole
crickets to inoculative concentrations of nematodes on each participating golf course.
Infection stations consisted of two sound traps each consisting of 1) an emitter, 2) a
fiberglass funnel, approximately 1.4 m in diameter and suspended from a metal yoke
attached to 10 cm x 10 cm wooden support posts, and 3) infective-stage nematodes held
beneath the funnel in a bucket or applied directly to the soil beneath the funnel. The
emitters, attached to the metal support yokes, sat directly over the funnels. Fig. 1
shows an infection station at a participating golf course.
The infection stations were operated during the spring mole cricket flight season.
The timing varies latitudinally in Florida, but generally occurs from February to mid-
or late June. Participants began operation of the infection stations when nematodes
were received and ended operation when mole cricket flight activity ceased.
In 1990 delivery of nematodes began 5 February for the courses located farthest
south (those in Collier, Palm Beach, and Lee counties). These courses received one
million nematodes each per shipment and received two subsequent shipments in mid-
April and May. The remaining courses received two shipments each of one and a half


March, 1993










Insect Behavioral Ecology '92 Parkman & Frank 77

























Fig. 1. An infection station on a participating golf course. The mercury vapor light
was added by golf course personnel at this location only in an attempt to enhance trap
attractiveness.



million nematodes; initial deliveries were made from mid-February to mid-March and
final deliveries were made from mid-April to early May. The courses farthest south
received three deliveries because their flight season is longer; therefore, viable
nematodes were required for a longer period of time.
In 1991, eight additional participants (seven golf courses and a sod farm) were
supplied with nematodes. Seven received their first shipments in mid-April and one
received its first shipment in mid-May. All but one received two shipments of one and
a half million nematodes each. Also, 13 of the original 21 participants received two
shipments in spring 1991, each consisting of one and a half million nematodes. Fig. 2
identifies participants and their locations.
Participants were given a choice of three methods by which to expose mole crickets
attracted to the emitters to S. scapterisci. Infective-stage nematodes could be 1) held
in moist sand in a 13.2 liter plastic bucket beneath the trap funnel, 2) held in damp foam
(common upholstery foam acquired from a local fabric store) which was cut into 2 cm x
2 cm x 1.3 cm cubes, the cubes held in plastic buckets beneath the funnels, or 3) applied
directly to the soil beneath the funnels (Fig. 3). The station depicted in Fig. 1 used the
first method. (A station using damp foam would appear the same except for the contents
of the buckets.)
For stations using methods one and two, mole crickets attracted to the traps and
landing in the funnels were collected into the buckets through a wire mesh tube. Within
the buckets they were exposed to nematodes for approximately one day. Finally, they
were released onto areas chosen by participants operating the stations. Any dead mole
crickets were placed approximately 2.5 to 3 cm beneath the soil surface. Mole crickets
attracted to an infection station using the third method fell through the open bottoms









Florida Entomologist 76(1)


. SHORES


Fig. 2. Location of 28 golf courses and one sod farm which made inoculative releases
of S. scapterisci. Participants whose names are in italics began releases in 1991. All
others began releases in 1990. See Table 1 for method of release used by each partici-
pant.


of the funnels, to be exposed to nematodes in the soil below. Mole crickets which
dispersed rapidly to other locations (Ngo & Beck 1982) would die, releasing a magnified
number of infective-stage nematodes. Mole crickets dying in the soil below the funnels
would maintain the nematode inoculum of the soil near the infection stations.
In autumn, the infection stations were used to trap flying mole crickets of the sub-
sequent generation, to detect infection by S. scapterisci and to determine the proportion
infected. Mole crickets attracted to the stations were trapped beneath the funnels in
buckets containing moist sand only. Individuals thus trapped were sieved from the
sand, placed individually into vials, and chilled before and during shipment to our labor-
atory in Gainesville. Numbers shipped were recorded and individuals were held for
observation. Those dying within 10 days of capture were held for nematode emergence.
If no nematodes emerged from a cadaver, it was dissected to determine the presence
of nematodes. All nematodes emerging from, or found within, dead mole crickets were
identified.

RESULTS AND DISCUSSION

Of the 21 golf courses which made releases of S. scapterisci in spring and summer
1990, 18 collected and sent mole crickets for evaluation the following autumn.


March, 1993









Insect Behavioral Ecology '92 Parkman & Frank


Nematode-infected mole crickets were collected at seven of these courses (Table 1). Of
the courses trapping infected individuals, six used the method of exposing nematodes
in sand, while one exposed them in foam. In autumn 1991, seven of these original
participants collected and sent mole crickets for evaluation. Of these only one, Pineview
Golf and Country Club, collected infected mole crickets. This same course collected
infected individuals in autumn 1990 (Table 1).
Six of the eight participants which began releases in spring 1991 collected and sent
mole crickets for evaluation the following autumn. Two of them trapped infected individ-
uals (Table 1). One exposed nematodes in sand and one exposed them in foam.
The collection of nematode-infected mole crickets several months after inoculative
releases into the previous generation of the pests indicates establishment of S. scap-
terisci. Establishment at only nine of the release sites is not disappointing because,
although several participants sent three or more shipments of mole crickets for evalua-
tion, many sent few or none (Table 1). Because many release sites were not surveyed
adequately, establishment of S. scapterisci may have gone undetected at some of them.
The method of exposing infective-stage nematodes in foam appeared to be more
successful in establishing S. scapterisci than the other two methods. Sixty-seven per-
cent (2 of 3) of the participants using the foam method and who sent mole crickets for
evaluation apparently established the nematode. Thirty-two percent (7 of 22) of those
who used the sand method and who sent mole crickets for evaluation appeared to
establish the nematode. However, because the foam method and that of applying
nematodes directly to the soil were not used by more participants and, therefore, were
not evaluated adequately, a valid comparison of the three methods could not be made.




SOUND TRAP
INFECTION METHODS





NEMATODES IN NEMATODES IN NEMATODES IN
BUCKETS OF SAND BUCKETS OF FOAM SOIL BENEATH TRAPS



EMATODES NEMATODE








NEMATODES


Fig. 3. The three methods by which Scapteriscus mole crickets were exposed to S.
scapterisci: nematodes held in buckets of moist sand, nematodes held in buckets of
damp foam, and nematodes applied directly to soil beneath the traps.










80 Florida Entomologist 76(1) March, 1993

TABLE 1.MOLE CRICKET COLLECTION OF PARTICIPANTS IN THE INOCULATIVE RE-
LEASE OF S. SCAPTERISCI AND RESULTS OF EVALUATION FOR ESTAB-
LISHMENT. EVALUATIONS MADE IN AUTUMN 1990 ANDd 1991.'

1990 1991

No. crickets2 % No. crickets2 %3
per shipment infection per shipment infection

Banyan (so) 21.3 (3) 0
Bay Hill (s)
Citrus Hills (s) 30.3 (3) 0 11 (1) 0
Countryside (s) 25.5 (2) 0 31.5 (2) 0
Cypress Creek (s) 7 (1) 28.6 (1)
Cypress Run (s) 36.8(4) 0
Delaire (s) 21.3 (3) 0
Fiddlesticks (s) 42.4 (5) 0 35.0 (2) 0
Foxfire (s) 31 (1) 0
Golden Hills (s)
Interlachen (s) 12 (1) 0
Northdale (s) 24 (1) 33.3(1) -
Pineview (s) 26.5 (2) 33.0 (2) 47 (1) 6.4 (1)
Quail Ridge (s) 9 (1) 0 34 (1) 0
Riviera (s) -- 4 (1) 0
(Volusia Co.)
Riviera (s) 45.2 (5) 16.9 (2)
(Collier Co.)
R. Poinciana (f) 35 (1) 2.9 (1)
Sun City (s) 49.0(3) 1.4 (1)
TPC (s) 43.5 (2) 2.8 (1)
Waterford (s) 29.5 (4) 0 22.5 (2) 0
Woodfield (s) 10 (1) 0
A. Duda Sod (s) 23 (1) 4.3(1)
Bobby Jones (s) 42.7(4) 0
Englewood (f) 22 (1) 0
Ft. Walton Bch. (s) 22.0(2) 0
Golf Hammock (s) 62.0(4) 0
Grand Cypress (f)
Oceans West (f) 20.5 (2) 9.1 (2)
Pompano Bch. (s)

'Letter in parentheses following a participant name indicates method of nematode exposure (s = sand, f = foam,
so soil beneath traps).
zMean number or number of crickets shipped and evaluated per shipment; number in parentheses is number of
shipments made.
"Mean percent or percent infection of crickets in shipments containing infected crickets. Number in parentheses
is number of shipments containing infected crickets.



The ideal conditions for successfully inoculating S. scapterisci into a Scapteriscus
population would be to expose large numbers of mole crickets to fresh infective-stage
nematodes. Based on conversations with participating golf course superintendents we
determined that ideal conditions occurred at several of the sites where S. scapterisci
was established, e.g., Northdale, Pineview, Riviera, Royal Poinciana, Sun City, and










Insect Behavioral Ecology '92 Parkman & Frank


Duda Sod. Ideal conditions had not occurred at Cypress Creek and Oceans West,
however, where S. scapterisci was established although relatively few mole cric-
kets were exposed soon after receipt of nematodes. Success under such less-than-ideal
conditions indicates that establishment is possible even when flying mole crickets are
not abundant.
The methods described above provide an economical use of this biological control
agent when infective-stage S. scapterisci are not adequately abundant or affordable for
broadcast application. Nematodes distributed at a few per m2 probably would not be
effective in view of numbers shown to be necessary to infect and kill one mole cricket
(Aguillera 1992). However, clusters of up to 50,000 juvenile nematodes, as might be
released by one dying mole cricket, would be sufficient to infect several mole crickets.
Our method of inoculative release, because it should result in clusters of nematodes at
sites to which mole crickets are attracted, may be expected to be more effective than
even distribution of an equal number of nematodes.
Sound-emitters currently are marketed by EcoSim (Gainesville, Florida) at a cost
of approximately $200 per unit. The cost of 3 million S. scapterisci from a commercial
supplier should be trivial because of development of mass-production methods (though
our low-volume production methods were labor-intensive, and thus expensive). Includ-
ing other equipment for infection station construction (approximately $140 for materials
per infection station; more for labor) and annual maintenance (perhaps 20% of cost of
the equipment), the cost would still be far less than that of broadcast use of S. scap-
terisci. Although the biopesticidal advantages of the nematode are lost by this method,
it should function to maintain an inoculum of nematodes at each site.


ACKNOWLEDGMENTS

We thank Robert Rehberg of the Florida Turfgrass Research Foundation for initial
inspiration for this project, Robert Yount of the Florida Turfgrass Association for his
receptiveness and support for it, and the superintendents of the 28 participating golf
courses and manager of one sod farm for their collaboration and financial support. T.
J. Walker supervised the design and production of the emitters. Nematodes for the
project were reared in 1990 by Marineide Aguillera with assistance of James Castner,
Suzette Broche, and Takuji Hayakawa, and in 1992 by one of us (JPP) with assistance
of Marineide Aguillera and Kurt Engle; those used in 1991 were donated by Biosys (a
biological pest control company) of Palo Alto, California. Identity of nematodes reared
from mole crickets collected by participants was confirmed by K. B. Nguyen. We thank
J. L. Capinera and T. J. Walker for reviews of a draft manuscript. This is University
of Florida, Institute of Food & Agricultural Sciences, journal series no. R-01408.


REFERENCES CITED

AGUILLERA, M. M. 1992. Steinernema scapterisci Nguyen & Smart, 1990: Bacterial
associates, culture, and pathogenicity. Ph. D. dissertation, Univ. Florida, vii
+ 132 p.
FRANK, J. H. 1990. Mole crickets and other arthropod pests of turf and pastures, pp.
131-139, in D. H. Habeck, F. D. Bennett, and J. H. Frank [eds.], Classical
biological control of insects and weeds in the southern United States. Southern
Co-op. Ser. Bull. 355: i-viii, 1-197.
HUDSON, W. G., J. H. FRANK, ANDJ. L. CASTNER. 1988. Biological control of mole
crickets (Orthoptera: Gryllotalpidae) in Florida. Ann. Entomol. Soc. America 34:
192-198.










82 Florida Entomologist 76(1) March, 1993

KAYA, H. K. 1990. Soil ecology, pp. 92-115 in R. Gaugler and H. K. Kaya [eds.].,
Entomopathogenic nematodes in biological control. CRC Press; Boca Raton.
NGO, D., AND BECK, H. W. 1982. Mark-release of sound-attracted mole crickets: Flight
behavior and implications for control. Florida Entomol. 65: 531-538.
NGUYEN, K. B., AND G. C. SMART. 1990. Steinernema scapterisci n. sp. (Rhabditida:
Steinernematidae). J. Nematol. 22: 187-199.
NGUYEN, K. B., AND G. C. SMART. 1991. Pathogenicity of Steinernema scapterisci to
selected invertebrates. J. Nematol. 23: 7-11.
PARKMAN, J. P., AND J. H. FRANK, 1992. Infection of sound-trapped mole crickets,
Scapteriscus spp., by Steinernema scapterisci. Florida Entomol. 75: 163-165.
PARKMAN, J. P., W. G. HUDSON, J. H. FRANK, K. B. NGUYEN, AND G. C. SMART.
Establishment and persistence of Steinernema scapterisci (Rhabditida: Steiner-
nematidae) in field populations of Scapteriscus spp. mole crickets (Orthoptera:
Gryllotalpidae). J. Entomol. Sci. (in press).
WALKER, T. J. 1982. Sound traps for sampling mole cricket flights (Orthoptera: Gryl-
lotalpidae: Scapteriscus). Florida Entomol. 65: 105-110.
WALKER, T. J. [ed.]. 1985. Mole crickets in Florida. Univ. Florida Agr. Exp. Stn. Bull.
846 (1984): i-iv, 1-54.






INTRODUCTION OF EXOTIC ENTOMOPATHOGENIC
NEMATODES (RHABDITIDA: HETERORHABDITIDAE AND
STEINERNEMATIDAE) FOR BIOLOGICAL CONTROL OF
INSECTS: POTENTIAL AND PROBLEMS

RICHARD K. JANSSON
Tropical Research and Education Center
Institute of Food and Agricultural Sciences
University of Florida
Homestead, Florida 33031

ABSTRACT

Interest in the use of entomopathogenic nematodes (Heterorhabditidae and Steiner-
nematidae) as biological control agents of insect pests has grown rapidly in recent years.
As a consequence, there has been an increase in the numbers of soil surveys for these
nematodes worldwide. These surveys have isolated large numbers of feral nematodes,
some of which are new species and strains. It is anticipated that considerable exchange
of nematode germplasm will occur between various laboratories. Some of these ex-
changes may result in the introduction of exotic nematodes into non-native lands. The
present paper reviews the potential and concomitant risks of nematode introductions
into non-native lands.

RESUME

En los fltimos afos se ha observado gran interns en el uso de los nematodos en-
tomopatog6nicos de las families Heterorhabditidae y Steinernematidae para el control
de insects plaga. Como consecuencia, se ha incrementado en todo el mundo el ndmero






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Insect BehavioralEcology-'92: Jansson


de muestreos de estos nemitodos. A trav6s de estos muestreos se han colectado muchos
nemAtodos de los cuales algunos son species desconocidas. Se prevee que va a occurrir
un gran intercambio de material gen6tico de nemAtodos entire laboratories, y algunos
de estos intercambios podrian resultar en la introducci6n de nemAtodos ex6ticos de un
area a otra. En este trabajo se revisa el potential y los riesgos de introducciones de
nemAtodos a areas ex6ticas.



Interest in the use of entomopathogenic nematodes for biological control of insect
pests is growing rapidly, largely because of the many attributes of these nematodes as
biological control agents, including their broad host range, high virulence and reproduc-
tive rates, ability to seek out and kill cryptic hosts, quick killing potential within 24-48
h, ease of application, reported safety to nontarget organisms, and improvements in
their mass-production, storage, and development of commercial products. This interest
led to a sudden wave of soil surveys for these nematodes in England (Hominick &
Briscoe 1990), Ireland (Griffin et al. 1991), Germany and Italy (Ehlers et al. 1991),
Finland (Vanninen et al. 1989), Hawaii (Hara et al. 1991), New Jersey (Gaugler et al.
1992b), and the Caribbean (R.K.J. et al., unpublished). In addition, earlier surveys
were conducted in Australia (Akhurst & Bedding 1986), Czechoslovakia (MrBaek 1980),
North Carolina (Akhurst & Brooks 1984), Florida (Beavers et al. 1983), Puerto Rico
(Roman & Beavers 1982), China, and the Netherlands (cited in Akhurst & Bedding
1986). These surveys have expanded the nematode germplasm available for research
and exchange which might increase the likelihood of introducing exotic nematodes into
non-native lands. There have been numerous introductions of foreign nematodes into
non-native lands. For example, Steinernema feltiae (Filipjev) is the most widely used
foreign nematode in the United States (R. Georgis, pers. comm.). In addition, S. car-
pocapsae (Weiser) All strain (from the United States) has been introduced into most
countries in Europe, except the United Kingdom (R. Georgis, pers. comm.). This paper
discusses the potential for introducing exotic nematodes for biological control of insect
pests. Specifically, factors that may enhance the likelihood of success of a release are
reviewed. Also, the risks of introducing exotic nematodes into non-native lands are
defined. It was not the intent of this paper to provide a complete review of factors that
limit establishment of nematodes nor those that might pose environmental risks. More
complete information on some of these topics is available elsewhere (Poinar 1979,
Gaugler & Kaya 1990, Howarth 1991).

PREDICTABLE BIOLOGICAL CONTROL WITH NEMATODES

Ehler (1990) identified predictability as the most intellectually challenging issue in
biological control. Entomopathogenic nematode research is no exception. Gaugler (1988)
noted that we need a more complete understanding of nematode interactions with en-
vironmental factors that may determine the outcome of nematode releases. Information
is lacking on the fate of nematodes introduced into the soil, on factors regulating their
population dynamics, on optimal environmental conditions that enhance development of
epizootics, on ecological barriers to infection, and on nematode interactions with biotic
factors in the soil (Gaugler 1988). Georgis & Gaugler (1991) noted that most failures of
entomopathogenic nematodes against Japanese beetle, Popillia japonica Newman,
grubs were due to the use of an unsuitable nematode strain or environmental conditions.
Gaugler (1993) noted that poor field results from releases of genetically improved
nematodes were attributable to their poor adaptation to the target host.
Goals established for the release of exotic nematodes are similar to those for release









84 Florida Entomologist 76(1) March, 1993

of native nematodes and may vary depending upon the target host and habitat. Gener-
ally, however, in agricultural environments, the aim of releases is to reduce and main-
tain pest populations to noneconomic levels for the duration of the crop. The success of
an introduction can be measured in two ways: the effect on the target pest population
and crop damage, and persistence in the target habitat. We need to increase our ability
to predict the outcome of the release to increase the likelihood of success with exotic
nematodes. These goals might be achieved by evaluating the potential effect of a variety
of biotic and abiotic factors, and the release strategy on suppression of the target host
population and establishment of the nematode.

FACTORS LIMITING SUCCESS OF INTRODUCED NEMATODES

Target Host and Habitat

The success of an introduction will depend upon the biology and behavior of the
target host (Kaya 1985). The stage of the target host is of central importance in the
nematode-host encounter. It is well known that certain life stages (e.g., larva) are more
susceptible to these nematodes than others (e.g., pupa and adult). Success of introduc-
tions will also be limited by the availability of the host at the time of the introduction.
This is especially true for these nematodes because of their short dispersal capability
in soil and their short-lived persistence. Hence, an adequate and constant supply of
hosts is needed to enhance recycling at the site of introduction. In addition, the be-
havioral, physiological, and morphological defensive mechanisms of the target host may
affect the potential of these nematodes as biological control agents and the outcome of
a release (Gaugler 1988, Kaya 1990).
The habitat of the target host is especially critical in any introduction program. In
general, nematode releases have been more successful against below-ground target
hosts or hosts in cryptic environments (Begley 1990, Klein 1990) than against above-
ground targets on foliage (Kaya 1985, 1986, Begley 1990). This is due primarily to the
short persistence of these nematodes on foliage because of the deleterious effects of
ultraviolet radiation (Gaugler & Boush 1978, Gaugler et al. 1992a), and their vulnerabil-
ity to desiccation and high temperatures (Kaya 1985, Begley 1990 and references
therein). However, the limited success of these nematodes against above-ground targets
should not dissuade research activity in this area. Innovative approaches and new
technologies may help to improve control against above-ground pests (Begley 1990).

Nematode Suitability and Quality

Another important factor that must be considered in any release program with these
nematodes is the selection of the candidate nematode for release. Georgis & Gaugler
(1991) found that many control failures in the field were due to the use of an unsuitable
nematode. It is well known that nematode strains vary in their virulence and infectivity
to a target host (Bedding et al. 1983, Molyneaux et al. 1983, Mannion 1992, Mannion &
Jansson 1992). For this reason, pre-release studies should be conducted in the laboratory
to identify the most virulent nematode(s) for a target host and associated habitat. It
should be noted, however, that laboratory results do not always agree with field results.
For example, the use of Petri plate assays to screen nematodes and assess virulence is
not recommended because it favors nematodes that nictate (Bedding 1990). Mannion &
Jansson (1992) found that laboratory results from Petri plate assays concurred with field
results (Jansson et al. 1990, 1992, 1993). However, a variety of bioassays, some of which
closely resemble the natural environment, should be conducted to select candidate
nematodes for field studies (Mannion 1992).










Insect BehavioralEcology-'92: Jansson


Certain quality control factors need to be addressed to enhance success of the release
once a candidate nematode has been selected. Gaugler & Georgis (1992) showed that
the method used to mass-produce nematodes had a significant effect on their efficacy
in field releases against Japanese beetle. Mass-production methodology was especially
important for heterorhabditid nematodes which were shown to be more efficacious when
reared in vivo or in solid media than when reared in liquid culture. In addition, the
length of time that these nematodes have been in storage before release may have a
significant effect on the outcome of the introduction (Westerman 1992). Older nematodes
with depleted lipid reserves have decreased motility and infectivity which undoubtedly
decreases field efficacy.

Seasonality

Climatic conditions during the experiment will affect the success of any introduction
and our ability to predict its outcome. Several studies showed that certain nematodes
perform better within a certain temperature regime. Conditions outside the optimum
range drastically affect the infectivity and efficacy of these nematodes. For example,
Kung et al. (1991) showed that the temperate nematode S. carpocapsae survived better
at temperatures between 50 and 250C than at 35C, whereas S. glaseri (Steiner), a
subtropical/tropical nematode (Poinar 1990), survived better at warmer temperatures.
Burman & Pye (1980) showed that S. carpocapsae displayed a thermal preference for
the temperature at which it was reared; infective juveniles actively moved through a
temperature gradient towards the temperature that they were acclimated to when
cultured. Kung et al. (1991) suggested that persistence of steinernematid nematodes is
enhanced in environments similar to the nematode's climatic origins. Thus, the species
and strain of nematode, origin, and its thermal preference will probably affect the
predictability of success of a release in a given climatic zone and season of the year. In
addition to temperature, precipitation patterns after release, which affect levels of
available soil moisture, will have a significant effect on the outcome of a release, espe-
cially when precipitation is sporadic and deficient and water holding capacity of soils is
poor.

Application Strategy

Application strategy is a complex factor that can significantly affect the success of
nematode releases. In planning a release, researchers need to consider the variety of
options: inundative vs. inoculative; periodicity of release; single species or strain release
vs. multiple species/strain release; time and placement of applicationss; application
methodology; and post-application methods to maximize efficacy, motility, and persis-
tence of nematodes. Several of these release strategies were discussed previously (Ehler
1990).
Most field studies have used inundative releases (Georgis 1990), and in most cases,
no clear release densities have been recommended. For inundative biological control,
the aim is to release a density that will maximize short-term biological control (Ehler
1990). The inconsistency of field trials and poor predictability have hampered our ability
to develop release levels. The numbers to release will undoubtedly be affected by sev-
eral factors, including the numbers of species and/or strains released, the climatic origin
of the candidate nematode(s), the quality of the nematode, the biology and behavior of
the target host and the nematode, and environmental conditions.
Inoculative releases aim to establish a biological control agent permanently in the
target habitat and some studies with nematodes suggested that this strategy has merit
(Parkman & Frank 1993, Gaugler et al. 1992b). The success of establishment is depen-









Florida Entomologist 76(1)


dent upon many of those factors that will limit the success of inundative releases plus
the availability of alternate hosts during periods of low host density, a high host-seeking
ability of the candidate nematode, and a high level of virulence and recycling.
Another release strategy that might affect the likelihood of success is the periodicity
of release. Jansson et al. (1991) showed that a single release of Heterorhabditis bac-
teriophora Poinar was adequate for controlling populations of the sweetpotato weevil,
Cylas formicarius(Fabricius). Parkman et al. (1993) also showed that a single release
was adequate for establishing S. scapterisci Nguyen & Smart for control of mole cric-
kets, Scapteriscus spp., in northern Florida. Other studies, however, found that multi-
ple releases may be needed (K. Smith, pers. comm.). In general, the periodicity of
release needed will vary depending upon the candidate nematode, the target host and
habitat, and environmental conditions (Jansson et al. 1991).
The number of species or strains to be released may also affect the likelihood for
success. Alatorre-Rosas & Kaya (1990, 1991) showed that steinernematid and
heterorhabditid nematodes compete for hosts. Heterorhabditid nematodes had a com-
petitive advantage when hosts were placed at a greater distance from the inoculation
site (Alatorre-Rosas & Kaya 1990). However, when hosts were close to the inoculation
site, steinernematids outcompeted heterorhabditids. Steinernematids also held a com-
petitive advantage over heterorhabditids in Petri plates even when heterorhabditids
were introduced 10 hours before steinernematids (Alatorre-Rosas & Kaya 1991). Total
mortality was greater when the two nematodes were applied in combination than when
the two were applied alone (Alatorre-Rosas & Kaya 1991). To my knowledge, no field
studies have been conducted to test their results. Entomopathogenic nematodes have
the potential to enable researchers to test this biological control theory in the field
(Ehler 1990).
Other factors that will affect the likelihood of success with exotic nematodes include
the method of application (manual vs. mechanical applications), time and placement of
applications, and post-application methods, such as irrigating, that help to maximize
dispersal and persistence of nematodes. Most of these have been discussed previously
(Kaya 1985, Gaugler 1988).

Soil Characteristics

Soil characteristics have a significant effect on the survivorship, motility, and infec-
tivity of entomopathogenic nematodes (Kaya 1985, 1990 and references therein, Barber-
check 1992). Wallace (1963) stated that the most important factors affecting nematodes
are pore size, moisture, oxygen (aeration), temperature, and chemistry of soil solution
(pH). Recent studies confirm that persistence and infectivity of these nematodes are
limited by soil type (Kung et al. 1990), soil temperature, moisture, ambient relative
humidity (Kung et al. 1991). The reader is referred to Kaya (1990) and Barbercheck
(1992) for a complete review on the effects of soil ecology on these nematodes. It is
important to note, however, that not all entomopathogenic nematodes are affected
equally in a given soil environment. Therefore, it is important to know the relationships
between survivorship, motility, and infection of the candidate nematode and the target
soil environment before release. Matching the appropriate nematode with a compatible
soil environment will enhance the likelihood of obtaining a successful introduction.

Compatibility with Pesticides

Nematode compatibility with pesticides is a neglected area of research. Because of
the diversity of pest complexes that attack most crops, it is likely that an introduced
nematode will encounter one or more pesticides. Many biological control programs that


March, 1993










Insect BehavioralEcology-'92: Jansson


rely on predators and/or parasitoids have been hampered by the lack of compatibility
between pesticides and introduced enemies. Compatibility of pesticides with nematodes
was studied in turfgrass (Zimmerman & Cranshaw 1990). They found that nematode
survivorship in various pesticides differed among nematode species. Surjusingh et al.
(1991) also found that tolerance to pesticides differed among nematode species. In gen-
eral, steinernematids were found to be more tolerant to pesticides than were
heterorhabditids. Gaugler & Campbell (1991) found that oxamyl was not compatible
with S. carpocapsae nor with H. bacteriophora. Kaya (1985) listed several other studies
demonstrating compatibility or incompatibility of nematodes and certain pesticides. No
generalizations on nematode compatibility with pesticides can be made at this time;
thus, the pesticides that a nematode might encounter, and their potential effect on the
nematode, should be known before a release to increase the likelihood of success.


Biotic Factors in Soil

Several studies demonstrated that biotic antagonists in the soil can affect the biolog-
ical control potential of a nematode significantly (Kaya 1990). These nematodes are
susceptible to infection by nematophagous fungi (Timper & Kaya 1992, Timper et al.
1991) and microsporidans (Kaya 1990); they can be caught by nematode-trapping fungi
(Kaya 1990), and preyed upon by mononchid and dorylaimid nematodes (Kaya 1990),
various mites (Epsky et al. 1988), and some collembolans (Epsky et al. 1988) and tardi-
grades (Kaya 1990 and references therein). In addition to antagonists, the outcome of
released nematodes might also be influenced by competitors in soil, such as fungi (Bar-
bercheck & Kaya 1990, 1991). They found that total mortality of beet armyworm,
Spodoptera exigua Hiibner, was greater when entomopathogenic nematodes were com-
bined with a fungus, Beauveria bassiana (Bals.) Vuill., than when each entomopathogen
was present alone; however, due to interspecific competition for hosts, the fungus re-
duced the level of mortality achieved by the nematode more than the nematode reduced
the mortality due to the fungus (Barbercheck & Kaya 1991).

RISKS OF INTRODUCING EXOTIC NEMATODES

As mentioned earlier, factors limiting the success of exotic nematodes are no differ-
ent from those that limit releases of native nematodes. However, because of the in-
creased nematode germplasm currently available from several surveys and the potential
for exchange and release of these nematodes, the potential environmental effect of
releases of exotic nematodes must be weighed carefully before release.
Howarth (1991) discussed environmental effects of classical biological control pro-
grams and indicated that certain biological parameters that might help or hinder an
exotic organism's risk potential need to be defined in order to weigh the risk of an
introduction. There are several risks associated with introducing exotic nematodes into
non-native lands. Some of these risks are similar to those described by Howarth (1991).
The most important factors that affect the level of risk for introducing these nematodes
include: the potential for range expansion; the host range of the candidate nematode
and the effect of its encounters with non-target organisms (including their threat to
human and animal health); the permanence of the introduction and the vulnerability of
the target habitat; the nematode's relationship with associated bacteria; and their effect
on community and ecosystem dynamics. It should be noted that any introduction that
results in establishment of nematodes will likely affect the environment (Ehler 1990).
Although Ehler (1990) restricted his criterion to only those introductions that result in
permanent establishment, in certain cases, especially when inundative releases might
be conducted in vulnerable habitats, a significant effect could occur even without a










Florida Entomologist 76(1)


permanent establishment of the nematode. Although it is generally accepted that these
nematodes pose little environmental risk, considerably more data are needed before any
generalizations can be made (Ehler 1990).

Range Expansion

The potential risk of an introduction might be affected by the nematode's ability to
expand its range and disperse beyond the site of introduction. This risk is difficult to
weigh because few studies have been conducted.
Active dispersal of this nematode in soil is well documented (Moyle & Kaya 1981,
Georgis & Poinar 1983a,b,c, Poinar & Hom 1986, Mannion & Jansson 1993). It is unlikely
that this mode of dispersal is a significant threat for range expansion because of the
limited distances that these nematodes can move. Mannion & Jansson (1993) showed
that a small percentage of G. mellonella larvae placed 60 cm from inoculation points
became infected up to 4 months after application suggesting that few nematodes dis-
persed 60 cm laterally from the inoculation site.
Passive movement of these nematodes poses a more serious threat for range expan-
sion. Movement of plant-parasitic nematodes on farm equipment, etc., is well known;
thus, similar modes of dispersal that move soil by humans might also help facilitate
movement of these nematodes. Several researchers speculated that entomopathogenic
nematodes could disperse beyond the infection site on mobile adult insects before adults
become infected (Glaser & Farrell 1935, Finney & Walker 1977). Timper et al. (1988)
showed that S. carpocapsae dispersed up to 11 m from the site of infection on infected
S. exigua adults. They suggested that this type of dispersal could help account for the
widespread dispersal of these nematodes locally and perhaps worldwide. Parkman &
Frank (1992) showed that newly-infected mole crickets dispersed the nematode S. scap-
terisci. Mobile larvae might also aid in dispersal before infection (Molyneux et al. 1983).
Epsky et al. (1988) showed that megostigmatid and oribatid mites could serve as phore-
tic hosts and help to disperse S. carpocapsae.

Host Range and Impact on Nontarget Organisms

Entomopathogenic nematodes are considered to have a broad insect host range
(Laumond et al. 1979, Poinar 1979, 1990, Gaugler 1981). However, in nature the host
range of these nematodes may be restricted to some extent by behavioral and ecological
barriers. For example, not all entomopathogenic nematodes have a broad host range.
Steinernema scapterisci is highly specific for mole cricket hosts, Scapteriscus spp.
(Nguyen & Smart 1991). Also, as mentioned earlier, virulence differs among nematodes
in a variety of insect hosts. Host range may also be restricted by susceptibility of the
life stage encountered by these nematodes after release. In addition, although many
insects are susceptible to these nematodes in the laboratory, not all are likely to be
infected in nature (Smart 1992). Also, although nematode-nontarget organisms encoun-
ters are inevitable, the result of these encounters will not be the same for all nematodes.
Several studies showed that these nematodes do not infect birds or mammals (Poinar
1990 and references therein). However, Kermarrec & Mauleon (1985) showed that these
nematodes have the potential to infect young toad tadpoles of Bufo marinus (L.). More
recently, Kermarrec et al. (1991) also showed that a lizard, Anolis marmoratus Dumbril
& Bibron, was affected adversely by some of these nematodes. Poinar Pnd Thomas
(1988) showed that S. carpocapsae and Heterorhabditis bacteriophora Poinar could kill
young tadpoles of Hyla regilla Baird & Girard and Xenopus laevis (Daudin).
Ishibashi & Kondo (1986) found that applications of steinernematid nematodes in-
creased native populations of Rhabditida and decreased populations of plant-parasitic


March, 1993









Insect BehavioralEcology-'92: Jansson 89

nematodes. Other research has shown that introduction of exotic nematodes may affect
populations of nontarget beneficial insects (Kaya 1984, 1986, Kaya et al. 1982, Laumond
et al. 1979).
Others found that these nematodes have little effect on nontarget invertebrates
(Poinar 1979 and references therein). Ishibashi et al. (1987) found that repeated inocu-
lation of soil with S. carpocapsae did not affect populations of collembolans and mites
adversely. Similar results were found by Klein & Georgis (1992). Georgis et al. (1991)
showed that applications of entomopathogenic nematodes did not affect the numbers of
nontarget invertebrates (Carabidae, Staphylinidae, Gryllidae, Histeridae, Collembola,
Gamasida, Actinedida, and Oribatida) adversely in the soil compared with applications
of chemical insecticides. However, in their study, all invertebrates were pooled by their
respective family and the effect of nematode releases on individual species of nontarget
invertebrates was not determined. Others showed that these nematodes did not ad-
versely affect populations of an earthworm (Capinera et al. 1982) and honey bees (Kaya
et al. 1982). Nickle et al. (1988) suggested that research on the effect of these nematodes
on nontarget invertebrates should be considered if an introduction of an exotic nematode
is planned in the United States. Despite the evidence, it is premature to conclude that
these nematodes are safe for nontarget invertebrates in all environments (Akhurst
1990). Too few long-term studies on the effects of these nematodes on the invertebrate
fauna in an environment have been conducted to draw any conclusions.

Effect on Community Dynamics and Displacement of Native Nematodes

Hominick & Reid (1990) raised the question: Do entomopathogenic nematodes influ-
ence the plant communities that exist in a given area through their effect on the soil
herbivores? A recent study showed that these nematodes were most common in the
U.K. in roadside verges, where there were mixed and diverse plant communities
(Hominick & Briscoe 1990). Insect herbivores affect early-stage succession (Brown &
Gange 1989). Thus, it is conceivable that these nematodes might exert an impact on
succession of communities by affecting soil herbivores (Hominick & Reid 1990). If these
nematodes play a significant role in successional changes in plant communities, then the
effect of an introduction would extend beyond nontarget organisms in soil communities
to entire ecosystems.
As mentioned earlier, nematode releases may affect populations of native plant-
parasitic and rhabditid nematodes (Ishibashi & Kondo 1986). A logical question one
might ask is: Can releases of exotic nematodes displace or replace populations of native
entomopathogenic nematodes? Displacement of native nematodes is an unknown risk,
but one that should be contemplated before releasing a newly discovered nematode
species into a non-native area. Pre-release studies on competition between native and
exotic nematodes for hosts would help to evaluate this potential risk.

Permanence of Introduction and Vulnerability of Target Habitat

Howarth (1991) noted that introduced enemies that persist for long periods of time
have a greater chance of affecting the environment adversely. Most field studies on
entomopathogenic nematodes have shown that these nematodes persist poorly.
Nematode persistence in soil is a function of host availability, host-seeking ability,
survivorship, reproduction, availability of alternate hosts, time and method of applica-
tion, quality of nematodes, and soil biotic and climatic factors (Kaya 1990 and references
therein). Recent studies, however, demonstrated that these nematodes have the ability
to persist in field soil well beyond the time of application and in some cases beyond the
duration of a suitable habitat. Jansson et al. (1991, 1993) showed that heterorhabditid










90 Florida Entomologist 76(1) March, 1993

nematodes, H. bacteriophora and Heterorhabditis sp., persisted for over 230 days after
application in sweet potato fields in southern Florida. Klein & Georgis (1992) found that
H. bacteriophora persisted through the winter in turfgrass plots for up to 290 days after
application. Parkman et al. (1993) showed that S. scapterisci persisted for over 5 years
after application in northern Florida.
Vulnerability of the target habitat was also listed as a factor that might affect the
level of risk in classical biological control programs (Howarth 1991). This is probably
also true for entomopathogenic nematodes. Island habitats have been reported to be
very vulnerable (Howarth 1991). Hominick (1991) suggested that the high incidence of
heterorhabditid nematodes near the coasts of many Caribbean and Pacific islands, as
well as in the U.K., might be due to their inadvertent movement by man in ship ballast.
The effect of these inadvertent introductions is not known, but is assumed to be minimal
because of the low incidence of these nematodes in other habitats on these islands (Hara
et al. 1990, Hominick & Briscoe 1991, Griffin et al. 1991, R.K.J. et al., unpublished).
The potential for these nematodes affecting the environment probably increases with a
decrease in latitude. Environmental conditions in tropical climates are more conducive
for nematode establishment due to more favorable soil, temperature, and precipitation
patterns, an abundance of hosts throughout the year, especially coleopterans and
lepidopterans which are very susceptible to these nematodes (Capinera & Epsky 1992).

Relationship with Bacteria

Another risk of introducing exotic nematodes might be their relationship with bac-
teria. This is especially true for previously unrecorded nematodes that were recently
isolated and for which there is no information on their symbiotic bacteria. Currently,
the taxonomy of Xenorhabdus, especially X. luminescens, is incomplete. Akhurst &
Boemare (1990) noted that X. luminescens, the bacterium associated with Heterorhab-
ditis, should probably be divided into several taxonomic groups. Similarly, the
taxonomic relationships between the bacteria and the Steinernematidae are not com-
pletely clear (Akhurst & Boemare 1990). Because at least one species of Xenorhabdus
(reported as X. luminescens [DNA hybridization group 5] but known to be a new
species [Akhurst & Boemare 1990]) was isolated from human wounds (Farmer et al.
1989), the symbionts of newly discovered nematodes may complicate government insec-
ticide registration in some countries. However, in reality, the symbiotic bacterium
probably poses little danger to the environment because it is not found normally outside
the host or the nematode. It was suggested that release of foreign nematodes into the
United States should not be considered until the nematode and its bacterial association
are known (Coulson et al. 1991).

SUMMARY AND FUTURE OUTLOOK

Introductions of exotic entomopathogenic nematodes may have potential for biolog-
ical control of insects in non-native lands. As shown by Bedding & Akhurst (1974),
classical biological control can be successful with nematodes. They showed that a siricid
wood wasp could be controlled with releases of the exotic nematode Beddingia (=
Deladenus) siricidicola Blinova & Korenchenko with little effect on nontarget or-
ganisms. A more recent study also indicated that release of an exotic nematode, S.
scapterisci, from South America against mole crickets, Scapteriscus spp.,in Florida has
potential for controlling these insects (Parkman et al. 1993).
Because entomopathogenic nematodes have a limited ability to expand their habitat
range once introduced, usually persist for short periods of time in most habitats, report-
edly have little affect on nontarget organisms, and pose a low risk to man and animals,









Insect BehavioralEcology-'92: Jansson


introductions of exotic nematodes should be encouraged. However, because we know
virtually nothing about recently isolated new species of Heterorhabditis and
Steinernema and their associated bacteria, release of these nematodes should await the
results of more rigorous pre-release studies on identification, host range, affect on
nontarget organisms, etc. Guidelines for importation, movement, and release of exotic
entomopathogenic nematodes in the United States are available (Nickle et al. 1988,
Coulson et al. 1991). It should be noted, however, that many researchers studying these
nematodes advocate less restrictive guidelines (Parkman et al. 1992, Smart 1992) than
those proposed by Nickle et al. (1988) and Coulson et al. (1991). It is important to
emphasize with regard to all introductions of exotic nematodes that the success of an
introduction will depend upon all of the biotic and abiotic factors described above. Thus,
researchers should consider these factors very carefully when planning releases of exotic
nematodes so that we can better predict the outcome.


ACKNOWLEDGMENTS

I thank M. Klein, R. Georgis, and R. Giblin-Davis for their criticisms and sugges-
tions and J. H. Frank and E. D. McCoy for their comments and invitation to participate
in this symposium. This paper was developed through research supported by U.S.D.A.,
C.S.R.S., Tropical/Subtropical Agriculture Program, Grant Nos. 88-34135-3564 and 91-
34135-6134 (to R.K.J.) managed by the Caribbean Basin Administrative Group (CBAG).
This is Florida Agricultural Experiment Station Journal Series No. R-02955


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