FHTET-2005-05

September 2005

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND

NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL

OF INSECTS AND MITES

Forest Health Technology

Enterprise Team

TECHNOLOGY 

TRANSFER

Invasive Species

ANN E. HAJEK, MICHAEL L. MCMANUS, ITALO DELALIBERA JÚNIOR

FHTET U.S. Department 

of Agriculture

Forest

Service

Cornell

University
Federal Recycling Program

Printed on recycled paper.

The Forest Health Technology Enterprise Team (FHTET) was created in 1995 by the Deputy Chief for State and Private Forestry, USDA Forest Service, to 

develop and deliver technologies to protect and improve the health of American 

forests. This book was published by FHTET as part of the technology transfer 

series.

http://www.fs.fed.us/foresthealth/technology/

The use of trade, firm, or corporation names in this publication is for information only 

and does not constitute an endorsement by the U.S. Department of Agriculture.

The U.S. Department of Agriculture (USDA) prohibits discrimination in all its programs and activities 

on the basis of race, color, national origin, sex, religion, age, disability, political beliefs, sexual orientation, 

or marital or family status. (Not all prohibited bases apply to all programs.) Persons with disabilities 

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etc.) should contact USDA’s TARGET Center at 202-720-2600 (voice and TDD).

To file a complaint of discrimination, write USDA, Director, Office of Civil Rights, Room 326-W, 

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202-720-5964 (voice and TDD). USDA is an equal opportunity provider and employer.

Cover photo by

Vince D’Amico

University of Delaware 

USDA-Forest Service Dept. Entomology 

Townsend Hall

Newark, DE 19716

610-368-4289

vdamico@fs.fed.us
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND

NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL

OF INSECTS AND MITES

ANN E. HAJEK

 DEPARTMENT OF ENTOMOLOGY, CORNELL UNIVERSITY

ITHACA, NEW YORK USA

MICHAEL L. MCMANUS

 USDA FOREST SERVICE, NORTHEASTERN RESEARCH STATION

HAMDEN, CONNECTICUT, USA

ITALO DELALIBERA JÚNIOR

Department of Entomology, Plant Pathology, and Agricultural Zoology

ESALQ-University of São Paulo

Piracicaba, Brazil 

For additional copies of this publication contact:

Richard Reardon

FHTET, USDA Forest Service

180 Canfield St.

Morgantown, WV 26505

U.S.A.

(304) 285-1566

rreardon@fs.fed.us
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

This publication would not have been possible without the help of many people including 

P. Allsopp, T. Andreadis, J. Becnel, A. Bellotti, G. Blissard, D. Boucias, G. Carner, A. Diss, 

J. Eilenberg, H. Evans, B. Federici, J. Fuxa, P. Gullan, I. Hall, L. Harrington, D. Haugen, E. 

R. Hoebeke, K. Hodge, J. Huber, R. Humber, C. Ignoffo, P. Klasmer, M. Klein, C. Lange, J. 

Liebherr, J. Martin, S. Merkel, M. Montgomery, D. Moore, C. Nielsen, D. Pilarska, J. Podgwaite, 

G. Poinar, B. Ratcliffe, M. Schlabach, A. Sharov, D. Smith, L. Smith, L. Solter, D. Sosa-Gomez, P. 

Stock, D. Streett, K. Teramoto, F. Vega, L. Volkman, J. Weiser, S. Woods, O. Zethner, S. Zinder, 

Yang Zhong-qi and many more. We also sincerely thank the excellent Entomology and Mann 

Libraries at Cornell University and the helpful and very knowledgeable librarians who work 

there.

 Finally, we wish to thank Dick Reardon, USDA Forest Service, Forest Health Technology 

Enterprise Team (FHTET), for supporting this project, and Chuck Benedict, USDA Forest 

Service, FHTET/ITX, for editing, designing, and laying out this publication, and for sheparding 

it through production.

Acknolwedgements
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

Table of Contents

Introduction........................................................................................................1

Table A: Exotic viruses released and target hosts................................................3

Table B: Exotic bacteria released and target hosts............................................11

Table C: Exotic fungi and oomycetes released and target hosts........................13

Table D: Exotic microsporidia released and target hosts...................................30

Table E: Exotic nematodes released and target hosts........................................32

Table F: Accidental introductions of pathogens and nematodes........................39

Chart 1: Classification of pathogens and nematodes purposefully introduced for 

classical biological control of insects and mites and accidentally introduced 

pathogens and nematodes infecting insects and mites......................................43

Chart 2: Classification of insects and mites targeted by classical biological 

control programs using pathogens and nematodes, and insect and mite hosts

 of accidentally introduced pathogens and nematodes....................................46

References........................................................................................................49

Indexes.............................................................................................................56

 Scientific names of insect and mite hosts....................................................................56

 Families of insect and mite hosts.................................................................................57

 Scientific names of pathogens and nematodes............................................................57

 Higher classification of pathogens and nematodes...............................................................58

 Release countries and regions.....................................................................................58

 Source countries and regions.....................................................................................59

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1

Introduction

Classical biological control is a strategy that has been defined as “The intentional introduction of an exotic biological control agent for permanent establishment and long-term pest control” (41). Numerous summaries of the many classical biological control programs 

have been published (e.g., 28, 55, 74). This strategy has 

been used extensively to control weeds and arthropods 

pests. For control of weeds phytophagous arthropods 

have principally been used and for control of arthropod 

pests parasitoids and predators have principally been 

used (58). 

Most programs using pathogens and nematodes 

for control of insects and mites have focused on mass 

production and inundative release. As long-term solutions for insect and mite pests (i.e., use in classical biological control programs), pathogens and nematodes 

have been used much less frequently when compared 

with parasitoids and predators (46, 59). Interestingly, 

while some classical biological control programs using 

pathogens and nematodes have been very successful in 

controlling insect and mite pests, some accidental introductions of entomopathogenic agents have also yielded 

substantial and long-term control.

We believe this publication is the most complete 

catalogue to date of classical biological control programs 

that have used pathogens and nematodes to control arthropod pests. It was difficult to find many of the classical biological control programs listed in the tables that 

follow; probably, we have not listed them all. Likewise, 

it was often difficult determining whether a release program should be included in this catalogue, particularly 

when a program was implemented many years ago and/

or not thoroughly documented. Thus, we used the following criteria for including programs in this catalogue:

1. The target pest was an insect or mite.

2. The microbial pathogen or nematode was an exotic (non-native) in the area of release. We include programs where the species of microbe or 

nematode was exotic (introduced) as well as programs where only the strain or biotype released 

was exotic.

3. Whether the releases were successful or not, the 

establishment of the microbe appeared to be a 

goal (i.e., long-term establishment and control) 

and was either investigated or discussed or, for 

older programs, we can infer that establishment 

of the pathogen or nematode was a goal of the 

program.

 (Note: Intentionally, we did not include examples 

of early widespread introductions of entomopathogens 

that were later shown to be questionably pathogenic, or 

widespread introductions where contaminants were actually released instead of the intended organisms [e.g., 

see 25, 147].)

Organization of tables and charts

Tables

Intentional and accidental releases of entomopathogens 

are grouped according to specific pathogen and nematode groups, and are presented in Tables A through F. 

Column headings and descriptions of contents are as 

follows:

Pest Group and Species

Only pestiferous insect and mite hosts are included. 

Taxonomic grouping, scientific names and synonyms 

for species names used in the publications cited or in the 

literature, are provided. If known, common names for 

pests are included.

Biological Control Agent

All natural enemies listed are exotic to their respective 

areas of release, i.e., either the species or the strain released was exotic, and include viruses, bacteria, fungi, 

a chromist, microsporidia and nematodes. Scientific 

names and synonyms are provided.
2 Introduction

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

Release date (year)

The year of release is listed, providing the intent of the 

release was to establish the pathogen or nematode in 

the release area. In some cases, pathogen or nematode 

levels declined over time, so agents were re-introduced. 

In other cases, pathogens have been re–introduced 

throughout a region over a period of years because the 

agents spread slowly on their own. In both cases, we list 

only the year or years of the initial releases; the dates of 

second or third introductions, or release in later years 

in the same general region, are included only if the initial release failed, or the pathogens used in subsequent 

releases were from a different source or sources. In the 

case of accidental introductions (Table F), the year the 

agent was first found is listed.

Release country, source and results from 

introduction

Releases are presented separately for geographically isolated areas and are listed by the country, or in a few instances by region (e.g. “Europe”), where the release was 

made (given in capital letters, e.g., BRAZIL). In some 

cases, a pathogen or nematode was released in more 

than one area within the same country. If release areas 

are isolated from one another, these introductions are 

considered separate introductions. For example, releases 

of Romanomermis culicivorax (originating from Louisiana) in both Maryland and California would be considered separate introductions although both are states 

within the same country because these release areas are 

geographically separated from one another and the sites 

differ in climate and topographies. The exception to 

this would be the release of a pathogen or nematode on 

proximate islands of the same country, e.g., in the many 

island groups in the south Pacific. If it appears that the 

introductions of pathogens or nematodes on proximate 

islands within a group were part of the same program, 

only the initial introduction is listed.

The geographical location where the pathogen or 

nematode was acquired for the release follows the release country (after Ex., e.g. Ex. China). Whenever appropriate, microbes from different source locations are 

listed separately. 

Results of introductions are provided as brief summaries of establishment, control, and persistence. We 

found that it is not always easy to classify control programs by strategy (i.e., classical biological control vs. 

inundative augmentation) and there are multitudes of 

programs where pathogens and nematodes have been 

released inundatively. For studies to be included in this 

catalogue, there must be some documented evidence 

that, whether the pathogen persisted or not after release, 

the intent of the program was to establish the pathogen 

in the release area for long-term, not temporary, control. 

Some older, poorly documented programs are exceptions and are included when we inferred the goal was 

establishment.

Clear summaries of results from introductions cannot always be found. In some cases, this is because not 

enough time has transpired since the release to see an 

effect. Unfortunately, in other cases, especially in earlier programs, we could find no documentation of what 

happened after releases.

Pest origin

I = Introduced (exotic)

N = Native (endemic)

? = Origin unknown

Charts

Chart 1 provides the classification for pathogens and 

nematodes included in the catalogue either as intentional 

or accidental releases. 

Chart 2 provides the classification for insect and mite 

hosts targeted by pathogens or nematodes that were either intentionally or accidentally introduced.

References

The reference list does not include every mention of a 

classical biological control introduction of a pathogen or 

nematode. Rather, it includes selected sources providing 

the information presented in this catalogue. If the information included in the catalogue has not been published, 

the individual providing the information is cited.
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

3

 ORDER: COLEOPTERA

 FAMILY

Scarabaeidae

 SPECIES

Oryctes rhinoceros (L.)

(rhinoceros beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Oryctes rhinoceros virus (OrV) (= 

Rhabdionvirus oryctes (Huger); 

= Baculovirus oryctes Huger) 

(Currently unassigned; previously 

in Baculoviridae)

1967 SAMOA (= WESTERN 

SAMOA) Ex. Malaysia. 

Established in 1 year and 

spread. Between 1973-75, 

adult infection decreased 

from 63 to 35% and although 

total population density also 

declined, damage was noticed 

again. Virus was re-released 

1975-1978 with a resulting 

decline in damage.

I 14, 89, 90, 145, 

163

1967 TOKELAU ISLANDS Ex. 

Malaysia. Released on 

Nukunonu Atoll. Established 

and by 1973 39% of beetles 

infected and only 1.5-6.5% of 

palm fronds damaged.

I 14, 145, 164

1970-74 FIJI Ex. Samoa (= Western 

Samoa). Established and 

by 1974 57-68% of beetles 

infected. Damage decreased 

significantly 12-18 months 

after virus establishment.

I 12, 13, 14, 145

1970 PALAU Ex. Samoa (= Western 

Samoa). Established on 

Babeldaob Isl., controlling 

beetles.

I 129, 145

1983 PALAU Ex. Samoa (= Western 

Samoa). Released on Peleliu 

Isl. and “other places where 

beetle problems were evident,” 

resulting in beetle control.

I 129

TABLE A: EXOTIC VIRUSES RELEASED AND TARGET HOSTS
4 Table A: Exotic viruses released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1970-71 WALLIS ISLAND Ex. 

Samoa (= Western Samoa). 

Established, < 2 months 

after release spread over 

entire island. In 1 year beetle 

populations decreased by 60-

80% and damage decreased 

by 82%. Average number 

infested palms reduced from 

60% in 1967 to 20% in 1981.

I 14, 56, 62, 145

1970-71 TONGA Ex. Samoa (= 

Western Samoa). Released 

in Tongatapu. Established, 

epizootics developed in 5 

months and virus spread at 

2-3 km/month, beetles and 

damage reduced. After 7 

years, 84% of adult beetles 

infected throughout population 

and damage remained low (< 

5% of palm crowns surveyed).

I 145, 160, 161

1970-72 MAURITIUS Ex. Samoa (= 

Western Samoa). Established, 

beetle populations declined 

sharply from 1970. At least 

through 1976-77, damage 

reduced by 60-95%.

I 14, 105

1972 AMERICAN SAMOA Ex. 

Samoa (= Western Samoa). 

Established, virus spread 0.8-

1.6 km/month and damage 

declined.

I 14, 145

1978-79 PAPUA NEW GUINEA Ex. 

Samoa (= Western Samoa). 

Released on 3 islands; 

established at nearly all sites, 

spread at 1 km/month.

I 53

1983-84 INDIA Ex. India (Kerala). 

Released on Minicoy Island. 

Established within 9 months, 

pest suppressed to low levels 

and damage reduced. Pest 

remained at low levels 3.5 

years after release.

N 101

COLEOPTERAScarabaeidaeOryctes rhinoceros (L.)Oryctes rhinoceros virus (OrV)
Table A: Exotic viruses released and target hosts 5

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1987 INDIA Ex. India (Kerala). 

Released at 4 locations on 

Andaman Islands. Palm 

damage reduced by 90% 

within 43 months of release, 

large reduction in numbers 

of adults and numbers of 

breeding sites. Virus spread 

at 1 km/year. By 1996, beetle 

populations remained at low 

levels.

 N 71

1984-85 MALDIVES Ex. Philippines, 

Tanzania & Malaysia. 

Established and caused highly 

significant reduction in palm 

damage on most islands 

where released. Different 

strains released and one strain 

(X2B) consistently yielded 

better infection and pest 

reduction.

N 31, 165

 SPECIES

Oryctes monoceros (Olivier)

(rhinoceros beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Oryctes rhinoceros virus (OrV) (= 

Rhabdionvirus oryctes (Huger); = 

Baculovirus oryctes Huger)

(Currently unassigned; previously 

in Baculoviridae)

1973 SEYCHELLES Ex. Samoa. 

Released on Mahé, Praslin 

Island group & La Digue. 

Establishment confirmed in 

1986 on Praslin Island group 

only, with infection 70-90%.

N 86, 87

1981-83 SEYCHELLES Ex. Praslin 

Island group. Established 

on Mahé & Ste. Anne with 

20-50% infection and 30% 

reduction in beetle population.

N 86, 87

1983-87 TANZANIA Ex. Philippines & 

Samoa (= Western Samoa). 

Established at 2 sites, with 40-

60% infection after 1-1.5 years 

but reduction in frond damage 

not sustained by 1988.

N 123, 131

COLEOPTERAScarabaeidaeOryctes rhinoceros (L.)
6 Table A: Exotic viruses released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Scapanes australis (Boisduval)

(rhinoceros beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Oryctes rhinoceros virus (OrV) (= 

Rhabdionvirus oryctes (Huger); = 

Baculovirus oryctes Huger)

(Currently unassigned; previously 

in Baculoviridae)

before 1979SOLOMON ISLANDS Ex. Fiji.

Virus released on Guadalcanal 

but fate unknown.

N 139

 ORDER: LEPIDOPTERA

 FAMILY

Zygaenidae

 SPECIES

Harrisina brillians (Barnes & McDunnough) 

 (western grapeleaf skeletonizer)

 BIOLOGICAL CONTROL AGENT Release 

date

Country, source, and results 

from introduction

Pest 

origin

References

Harrisina brillians Granulovirus 

(HbGV)

(Baculoviridae)

1981-1982 UNITED STATES OF 

AMERICA Ex. Mexico & 

USA (Arizona). Released 

in central California (Tulare 

Co.). Established, epizootics 

develop in high density host 

populations. Overall, lowers 

general equilibrium density of 

host populations.

I 144

 FAMILY

Lymantriidae

 SPECIES

Lymantria dispar (L.)

(gypsy moth)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Lymantria dispar Multicapsid 

Nucleopolyhedrovirus (LdMNPV)

(Baculoviridae)

1972 SARDINIA Ex. Serbia. 

Established, high levels of 

larval mortality year of release, 

> 40% infection the next year 

and spread over 300 hectares.

N 88

COLEOPTERAScarabaeidae
Table A: Exotic viruses released and target hosts 7

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Lymantria monacha (L.)

(nun moth)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Lymantria monacha

Nucleopolyhedrovirus

(Baculoviridae)

1973-4 DENMARK Ex. Sweden and 

West Germany. Released 

in Silkeborg (1973), 90% 

infection year of release and, 

in 1974, no serious defoliation 

within and directly around 

virus-release stands while 

insecticides had to be applied 

to other areas. In 1975, no 

virus was found in the few 

larvae collected. In Grindsted 

(1974), the population 

collapsed the year of virus 

release but it is suggested that 

other factors, including the 

native virus, played important 

parts.

N 167

 FAMILY

Noctuidae

 SPECIES

Anticarsia gemmatalis (Hübner)

(velvetbean caterpillar)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Anticarsia gemmatalis Multicapsid 

Nucleopolyhedrovirus (AgMNPV)

(Baculoviridae)

1979-80 UNITED STATES OF 

AMERICA Ex. Brazil (Santa 

Catarina). Released in South 

Carolina. 59-86% infection 

the season of release but no 

infections found 1 year after 

release.

N 8, 23

1990-91 UNITED STATES OF 

AMERICA Ex. Brazil. 

Released in soybean fields 

in Louisiana. Established, 

causing 25-100% infection 

the year of release and 4-49% 

infection for years 2-4 after 

release, even in rotated fields.

N 47

LEPIDOPTERA

Lymantriidae
8 Table A: Exotic viruses released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Trichoplusia ni (Hübner)

(cabbage looper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Trichoplusia ni

Nucleopolyhedrovirus (TnNPV)

(Baculoviridae)

1970 COLOMBIA Ex. USA 

(California). Persisted 

after release, controlling 

subsequent pest generations. 

I 15, 33

 SPECIES

Pseudoplusia includens (Walker)

(soybean looper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Pseudoplusia 

includens Singlecapsid 

Nucleopolyhedrovirus (PiSNPV)

(Baculoviridae)

1975-77 UNITED STATES OF 

AMERICA Ex. Guatemala. 

Released in soybean fields 

in Louisiana, established, 

38-63% infection 12-15 years 

after introduction. 

N 48

 SPECIES

Agrotis segetum (Denis & Schiffermüller)

(turnip moth)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Agrotis segetum Granulovirus 

(AsGV)

(Baculoviridae)

1975-80 DENMARK Ex. Austria. 

Released in Lammefjord. 

Caused 65-70% reduction in 

damage soon after release 

and thought to have spread 10 

m from release. One year after 

release, ca. 99% of infectivity 

of virus applied to soils had 

been lost.

N 166, 168, 171

LEPIDOPTERANoctuidae
Table A: Exotic viruses released and target hosts 9

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: HYMENOPTERA

 FAMILY

Diprionidae

 SPECIES

Neodiprion sertifer (Geoffrey)

 (European pine sawfly)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Neodiprion sertifer 

Nucleopolyhedrovirus (NeseNPV)

(Baculoviridae)

1950 CANADA Ex. Sweden. 

Released in southern Ontario, 

near Strathroy. > 90% mortality 

14 days after release and virus 

persisted. Widely distributed 

for release in pine plantations, 

e.g., one introduction in 1951 

controlled an infestation 

over 100 acres within 3 

years. After introduction, this 

virus replaced insecticides 

for controlling hosts and 

provided long-term control. 

Today, host is a minor pest of 

plantations and ornamentals 

but occasionally can increase 

locally as natural spread and 

effectiveness of the virus is 

much reduced at low host 

densities.

I 19, 34, 39, 92

1951-52 UNITED STATES OF 

AMERICA Ex. Canada. 

Released in New Jersey, 

established and spread (ca. 

300 m from individual trees 

after release). Provided 

complete control.

I 39
10 Table A: Exotic viruses released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1952 UNITED STATES OF 

AMERICA Ex. USA (New 

Jersey). Released in Illinois. 

By 19 days after treatment, 

82-100% control. In 1953, 

spread was up to 80 m from 

treated area. Excellent control 

achieved. Virus from Canada 

released in Indiana from 1953 

and reported as maintaining 

adequate control over several 

years through recurring 

epizootics after establishment. 

I 39, 130

1961 UNITED KINGDOM 

(SCOTLAND) Ex. Canada. 

85% of colonies had infected 

individuals 24 days after 

release, resulting in very 

good control. In 1962, found 

to persist in treated areas but 

minimal spread.

N 35, 126

 SPECIES

Gilpinia hercyniae (Hartig) (= Diprion hercyniae (Hartig))

(European spruce sawfly)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Gilpinia hercyniae

Nucleopolyhedrovirus (GhNPV)

(Baculoviridae)

1943-45 CANADA Ex. Canada 

(Mainland). Released in 

Newfoundland. Established 

and by 1946 reported as 

prevalent over considerable 

areas surrounding release 

areas. 

I 4, 92

1950 CANADA Ex. Canada (New 

Brunswick). Released in an 

isolated host population in 

Sault Ste. Marie, Ontario, 

160 km (100 miles) beyond 

western distribution of insect. 

Established and spread 

rapidly through infested area. 

Epizootics occurred yearly 

(1950-1959), hosts kept below 

economic damage level.

I 20

HYMENOPTERADiprionidaeNeodiprion sertifer (Geoffrey)Neodiprion sertifer 
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

11

 ORDER: COLEOPTERA

 FAMILY

Scarabaeidae

 SPECIES

Papuana huebneri (Halmahera)

(taro beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Paenibacillus popilliae (Dutky)

(Bacillaceae)

1976 KIRIBATI Ex. Papua New 

Guinea & Solomon Islands. 

Released on So. Tarawa. 

Isolate from Papua New 

Guinea (type A1) caused 

infections 1 year after release.

I 149

 SPECIES

Oryctes rhinoceros (L.)

(rhinoceros beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Paenibacillus popilliae (Dutky)

(Bacillaceae)

1951 PALAU Ex. USA (strain from 

Japanese beetle, Popillia 

japonica Newman). Not 

recovered after release.

I 145

1957 AMERICAN SAMOA Ex. USA 

(strain from Japanese beetle, 

Popillia japonica Newman). Not 

recovered after release.

I 145

 SPECIES

Schizonycha sp.

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Paenibacillus popilliae (Dutky)

(Bacillaceae)

1956 KENYA Ex. USA (A & B strains 

from Japanese beetle, Popillia 

japonica Newman). Not 

recovered after release.

N 54

TABLE B: EXOTIC BACTERIA RELEASED AND TARGET HOSTS
12 Table B: Exotic bacteria released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Cochliotis melolonthoides (Gerstaecker)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Paenibacillus popilliae (Dutky)

(Bacillaceae)

1968 TANZANIA Ex. Probably USA. 

Japanese beetle, Popillia 

japonica Newman. Seemed to 

become established but this is 

not certain due to presence of 

an indigenous milky disease. 

N 54

COLEOPTERAScarabaeidae
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

13

 ORDER: ORTHOPTERA

 FAMILY

Acrididae

 SPECIES

Phaulacridium vittatum (Sjöstedt)

(wingless grasshopper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Entomophaga grylli (Fresenius) 

Batko, pathotype I

(Zygomycetes: 

Entomophthoraceae)

1984 AUSTRALIA Ex. USA 

(Arizona). Released near 

Canberra. Epizootics did 

not occur and permanent 

establishment questioned, 

efficacy unlikely.

N 96

 SPECIES

Various species, including Melanoplus bivittatus (Say), Melanoplus sanguinipes (F.), Camnula 

pellucida Scudder

(twostriped grasshopper, migratory grasshopper, clearwinged grasshopper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Entomophaga grylli (Fresenius) 

Batko, pathotype III

(Zygomycetes: 

Entomophthoraceae)

1989-91 UNITED STATES OF 

AMERICA Ex. Australia. 

Isolate chosen based on 

biology, similarity of climates 

and ability to infect species 

in both Oedipodinae and 

Melanoplinae, but not 

Hesperotettix viridis (Scudder). 

Released in North Dakota. 

Populations of some species 

declined in 1991-92 with 23% 

infection in 1992 at < 1 km 

from release, 1.7% in 1993 

and no infection in 1994 when 

host populations were low. 

Long-term establishment 

questionable.

N 18, 24, 25

TABLE C: EXOTIC FUNGI RELEASED AND TARGET HOSTS
14 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Melanoplus sanguinipes (F.)

(migratory grasshopper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Entomophaga grylli (Fresenius) 

Batko, pathotype III

(Zygomycetes: Entomophthoraceae)

1990 UNITED STATES OF 

AMERICA Ex. Australia.

Released in Alaska. No 

establishment reported.

N 24, 124

 ORDER: HEMIPTERA

 FAMILY

Cercopidae

 SPECIES

Aeneolamia flavilatera (Ulrich)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

1944 GUYANA Ex. Trinidad. 

Introduced by releasing 

infected adult froghoppers. 

Established, considered 

unsuccessful for control 

but < 1 year later abundant 

infections ca. 32 km away. 

Unknown whether this was due 

to introduced or indigenous 

fungus.

N 29

 FAMILY

Cicadellidae

 SPECIES

Empoasca fabae (Harris)

(potato leafhopper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Zoophthora radicans (Brefeld) 

Batko

(Zygomycetes: 

Entomophthoraceae)

1984 UNITED STATES OF 

AMERICA Ex. Brazil. 

Introduced to Illinois. No 

establishment.

I 65, 93

ORTHOPTERAAcrididae
Table C: Exotic fungi and oomycetes released and target hosts 15

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Unspecified species

(leafhopper)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Unknown fungus 1906 UNITED STATES OF 

AMERICA Ex. Australia and 

Fiji, where they infected 

leafhopper eggs. Released 

in Hawaii. Establishment not 

confirmed.

I? 114

 FAMILY

Aphididae

 SPECIES

Therioaphis maculata (Buckton)

(spotted alfalfa aphid)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Zoophthora radicans (Brefeld) 

Batko

(Zygomycetes: 

Entomophthoraceae)

1979 AUSTRALIA Ex. Israel. Isolate 

chosen in part due to similar 

climate. Released in New 

South Wales. Became widely 

distributed in New South Wales 

and southern Queensland, 

causing epizootics in late 

summer/autumn; only the first 

aphid outbreaks in spring likely 

to escape infection.

I 97, 98, 99

HEMIPTERACicadellidae
16 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Aphis gossypii Glover

(cotton aphid)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Neozygites fresenii (Nowakowski) 

Batko

(Zygomycetes: Neozygitaceae)

1994-95 UNITED STATES OF 

AMERICA Ex. USA (Arkansas). 

Released in San Joaquin 

Valley, California. Cycling 

during release seasons with 

infection levels that would have 

initiated epizootics in Arkansas 

but epizootics did not occur. 

Persisted until end of release 

seasons but not recovered 

1997-2001, so long-term 

establishment questionable.

I 52, 142

 SPECIES

Metopolophium dirhodum (Walker), plus other cereal aphids

(rose-grain aphid)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Pandora neoaphidis (Remaudière 

& Hennebert) Humber

(Zygomycetes: 

Entomophthoraceae)

1982 BELGIUM Ex. Brazil. Isolate 

chosen due to good in vitro 

growth. Limited transmission 

in field after release, probably 

because few condia are 

produced by isolate. Limited 

transmission suggests poor 

chance of establishment.

N 85

HEMIPTERAAphididae
Table C: Exotic fungi and oomycetes released and target hosts 17

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Macrosiphum solanifolii (Ashmead)

(potato aphid)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Probably in Lecanicillium (=

Verticillium) lecanii species 

complex (Reported as 

Acrostalagmus sp.)

(Ascomycetes: Anamorph of 

Hypocreales)

1955 UNITED STATES OF 

AMERICA Ex. USA (Hawaii). 

Released in Maine. Diseased 

aphids found 3 weeks after 

release and one infected 

specimen found in 1958. 

Unknown if permanently 

established.

N 132, 133

 FAMILY

Aleyrodidae

 SPECIES

Singhiella citrifolii (Morgan) (= Dialeurodes citrifolii Morgan)

(cloudywinged whitefly)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Aschersonia goldiana Saccardo 

& Ellis

(Ascomycetes: Anamorph of 

Hypocreales)

1924 BERMUDA Ex. USA (Florida).

Considered established 

in 1925 but only provided 

efficient control in well-shaded 

situations.

I 109, 110

HEMIPTERAAphididae
18 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Dialeurodes citri (Ashmead)

(citrus whitefly)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Aschersonia spp.

(Ascomycetes: Anamorphs of 

Hypocreales)

1960-1964 GEORGIA Ex. 11 species and 

forms from China, Trinidad, 

Vietnam, Cuba, Japan & India. 

No information on whether all 

strains became established. 

The most aggressive was A. 

placenta Berkeley & Broome 

from Vietnam and China, 

giving up to 90% infection in 

Adzharia in favorable weather 

but was inhibited by drought. In 

1980-84, in areas where large 

complexes of natural enemies 

occurred in citrus plantations 

(including fungi in this genus), 

the pest was kept below the 

economic threshold. 

I 70, 120

1961-1964 AZERBAIJAN Ex. 11 species 

and forms from China, Trinidad, 

Vietnam, Cuba, Japan, USA 

& India. No information on 

whether all strains became 

established. About 80% larval 

mortality in citrus plantations 

under favorable conditions 

and fungus spread to new 

plantations. In 1980-84, in 

areas where large complexes 

of natural enemies occurred 

in citrus plantations (including 

fungi in this genus), the pest 

was kept below the economic 

threshold. 

I 70, 122

 SPECIES

Dialeurodes sp.

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Aschersonia aleyrodis Webber

(Ascomycetes: Anamorph of 

Hypocreales)

1926 BERMUDA Ex. USA (Florida).

Establishment and persistence 

not reported.

? 111

HEMIPTERAAleyrodidae
Table C: Exotic fungi and oomycetes released and target hosts 19

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Aleurodicus cocois (Curtis), Aleurothrixus floccosus (Maskell)

(coconut whitefly, woolly whitefly)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Aschersonia aleyrodis Webber

(Ascomycetes: Anamorph of 

Hypocreales)

before 1920VIRGIN ISLANDS Ex. 

Unknown. No establishment 

due to high winds and drought.

N? 154

 SPECIES

Unspecified whitefly species

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Unknown fungus 1909 UNITED STATES OF 

AMERICA Ex. USA (Florida). 

One species released in Hawaii 

against whitefly. Results not 

reported.

? 81

 FAMILY

Coccidae

 SPECIES

Coccus viridis (Green)

(green scale)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Unidentified fungus (possibly 

in Lecanicillium (= Verticillium) 

lecanii species complex)

(Ascomycetes: Anamorph of 

Hypocreales)

1928 or 

before

UNITED STATES OF 

AMERICA Ex. USA (Florida).

Released in Hawaii. 

Established and provided 

effective control.

I 69, 146

HEMIPTERAAleyrodidae
20 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Coccus viridis (Green), Eucalymnatus tessellatus (Signoret), Ceroplastes rubens (Maskell)

(green scale, tessellated scale, red wax scale)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Lecanicillium lecanii

(Zimmerman) Gams & Zare (= 

Verticillium lecanii (Zimmerman); 

= Cephalosporium lecanii 

Zimmerman)

(Ascomycetes: Anamorph of 

Hypocreales) 

1911 SEYCHELLES Ex. Sri Lanka. 

Established and largely 

controlled scale populations.

I 1

 SPECIES

Various species of lecaniine scales

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Lecanicillium lecanii

(Zimmerman) Gams & Zare (= 

Verticillium lecanii (Zimmerman); 

= Cephalosporium lecanii 

Zimmerman)

(Ascomycetes: Anamorph of 

Hypocreales)

before 1933SEYCHELLES Ex. India. 

Well established on lecaniine 

scales, especially Coccus 

viridis (Green) on coffee, and 

spread widely.

? 139

 SPECIES

Unspecified species of coccids

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Unknown fungi 1897 UNITED STATES OF 

AMERICA Ex. Unrecorded 

(Unknown). 2 species released 

in Hawaii against Coccidae. 

Establishment and spread over 

most parts of the islands.

? 79

HEMIPTERACoccidae
Table C: Exotic fungi and oomycetes released and target hosts 21

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 FAMILY

Diaspididae

 SPECIES

Aonidiella aurantii (Maskell)

(California red scale)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Fusarium coccophilum

(Desmazieres) Wollenweber 

& Reinking (= Fusarium 

episphaerea f. coccophila Tul.); 

teleomorph = Nectria flammea

(Tulasne & Tulasne) Dingley

(Ascomycetes: Anamorph of 

Hypocreales)

1900 ARGENTINA Ex. USA. 

Established and occasionally 

caused up to 90% mortality in 

northeastern and northwestern 

regions. 

I 32

 SPECIES

Cornuaspis beckii (Newman) (= Lepidosaphes beckii (Newman))

(purple scale, mussel scale)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Fusarium coccophilum

(Desmazieres) Wollenweber 

& Reinking (= Sphaerostilbe 

coccophila Tul.); teleomorph 

= Nectria flammea (Tulasne & 

Tulasne) Dingley

(Ascomycetes: Anamorph of 

Hypocreales)

1905 or 

before

UNITED STATES OF 

AMERICA Ex. USA (Florida). 

Introduced to Hawaii. 

Established and locally 

abundant but control only 

partial.

I 80

1926 BERMUDA Ex. USA (Florida). 

Establishment and persistence 

not recorded.

I 111

Podonectria coccicola Petch

(Ascomycetes: Tubeufiaceae)

1926 BERMUDA Ex. USA (Florida). 

Establishment and persistence 

not recorded.

I 111

Triblidium caespitosum Cooke 

& Masee (= Myrangium duriaei

Montagne & Berkeley)

(Ascomycetes: Triblidiaceae)

1926 BERMUDA Ex. USA (Florida). 

Establishment and persistence 

not recorded.

I 111

HEMIPTERA
22 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Aspidiotus destructor Signoret

(coconut scale)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Fusarium juruanum P. Hennings 

(= Pseudomicrocera henningsii

(Koord.) Petch)

(Ascomycetes: Anamorph of 

Hypocreales)

1929 SEYCHELLES Ex. Sierra 

Leone. Did not establish.

I 40

 SPECIES

Quadraspidiotus perniciosus (Comstock) (= Aspidiotus perniciosus (Comstock))

(San Jose scale)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Fusarium coccophilum

(Desmazieres) Wollenweber 

& Reinking (= Sphaerostilbe 

coccophila Tul.); teleomorph 

= Nectria flammea (Tulasne & 

Tulasne) Dingley

(Ascomycetes: Anamorph of 

Hypocreales)

1897 UNITED STATES OF 

AMERICA Ex. USA (Florida).

Released in California. As a 

result of this introduction, or 

a native fungus, scale nearly 

exterminated in southern 

California.

I 157

1897 UNITED STATES OF 

AMERICA Ex. USA (Florida).

Released in New Jersey. 

Established, overwintered, with 

abundant infection the following 

Sept. but this pathogen alone 

failed to provide adequate 

control.

I 134, 135

1898 UNITED STATES OF 

AMERICA Ex. USA (Florida). 

Released in Illinois by tying 

twigs with infected scales 

to trees. Overwintered and 

many scales infected but 

healthy scales still abundant. 

Hypothesized this fungus 

could add to effects of other 

natural enemies to provide 

a permanent check of scale 

populations but the level of 

fungus activity would depend 

on rainfall levels.

I 45

HEMIPTERADiaspididae
Table C: Exotic fungi and oomycetes released and target hosts 23

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: THYSANOPTERA

 FAMILY

Thripidae

 SPECIES

Thrips tabaci Lindeman

(onion thrips)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Neozygites parvispora (MacLeod 

& Carl) Remaudiére & Keller

(Zygomycetes: Neozygitaceae)

1973-76 BARBADOS Ex. Switzerland.

Released in onion field but no 

establishment.

I 29

 ORDER: COLEOPTERA

 FAMILY

Scarabaeidae

 SPECIES

Oryctes rhinoceros (L.)

(rhinoceros beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

1939 SAMOA (= WESTERN 

SAMOA) Ex. Java. This 

fungal species recovered after 

release but whether it was the 

introduced strain or a native 

strain is uncertain.

I 145

1952 WALLIS ISLAND Ex. Argentina. 

Results from release unknown.

I 145

1967 TOKELAU ISLANDS Ex. 

Samoa (= Western Samoa). 

Results from release unknown.

I 145

1969 TONGA Ex. Samoa (= Western 

Samoa). High levels of infection 

directly after release, infections 

still present 3 years later but 

prevalence extremely low.

I 145, 160
24 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Papuana huebneri (Halmahera)

(taro beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

1976 KIRIBATI Ex. Unknown. 

Released on southern Tarawa 

by Latch. Establishment not 

confirmed.

I 95

1995 KIRIBATI Ex. Papua New 

Guinea. Released on southern 

Tarawa. Persisted in soil 

through 2003, spread and 

exerted some control.

I 95, 149

 SPECIES

Phyllophaga smithi (Arrow) (= Lachnosterna smithi (Arrow); = Clemora smithi (Arrow); = Phytalus 

smithi (Arrow)

(sugar cane white grub)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Beauveria bassiana (Balsamo) 

Vuillemin (= Beauveria densa

(Link) Vuillemin)

(Ascomycetes: Anamorph of 

Hypocreales)

1932 MAURITIUS Ex. Unknown 

isolate from Imperial Bureau 

of Mycology, United Kingdom. 

Host population gradually 

declined and diseases may 

have played a part.

I 54, 102, 103, 

104

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

1932 MAURITIUS Ex. Unknown 

isolate from Imperial Bureau of 

Mycology, UK. Host population 

gradually declined and 

diseases may have played a 

part.

I 54, 102, 103, 

104

 SPECIES

Alissonotum impressicolle Arrow

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

1914 TAIWAN Ex. USA (Hawaii). 

Numbers of scarabs greatly 

reduced in fields where spores 

were released.

N 158, 159

COLEOPTERAScarabaeidae
Table C: Exotic fungi and oomycetes released and target hosts 25

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Dermolepida albohirtum (Waterhouse)

(greyback cane beetle)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

about 1914 AUSTRALIA Ex. Samoa. 

Released in Queensland but 

before release, had already 

been found infecting this host 

in Queensland. Post release, 

at times considerable numbers 

of grubs of intended host 

and Rhabdoscelus obscurus

(Boisduval) killed by this 

fungus.

N 155

 SPECIES

Lepidiota pruinosa Wied., Leucopholis irrorata Chevrolat 

(white grubs in sugar cane)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

1928 PHILIPPINES Ex. Australia 

(Queensland). Not effective 

control and “undoubtedly 

already present.”

N 125

 SPECIES

Various scarabs (including Anoplognathus sp., Lepidiota sp., etc.)

(white grubs in sugar cane)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Beauveria brongniartii (Saccardo) 

Petch (= Botrytis tenella Sacc.)

(Ascomycetes: Anamorph of 

Hypocreales)

1894-95 AUSTRALIA Ex. France. 

Releases in Queensland and 

New South Wales. Negative 

results in New South Wales 

after dissemination.

? 155

COLEOPTERAScarabaeidae
26 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Adoretus tenuimaculatus Waterhouse

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

before 1918FIJI Ex. Unknown. Some signs 

that this fungus acted as a 

check on the beetles.

I 78

 FAMILY

Curculionidae

 SPECIES

Sitona discoideus Gyllenhal

(sitona weevil)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Beauveria bassiana (Balsamo) 

Vuillemin

(Ascomycetes: Anamorph of 

Hypocreales)

1984 AUSTRALIA Ex. France 

(Montpellier). Released 

in southern Australia. No 

infections ever found.

I 3

 SPECIES

Otiorhynchus nodosus (Müller), Otiorhynchus arcticus (Fabricius)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Metarhizium anisopliae 

(Metschnikoff) Sorokin

(Ascomycetes: Anamorph of 

Hypocreales)

2003 ICELAND Ex. Faroe Islands 

(Havnardalur). Released in 

eroded areas in Haukadalur. 

Too early to determine effect.

N 108

COLEOPTERAScarabaeidae
Table C: Exotic fungi and oomycetes released and target hosts 27

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: DIPTERA

 FAMILY

Culicidae

 SPECIES

Aedes polynesiensis Marks

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Coelomomyces stegomyiae Keilin

(Chytridiomycetes: 

Coelomomycetaceae)

1958 TOKELAU ISLANDS Ex. 

Singapore. Released on 

Nukunonu Atoll. Established, 

by 1963 infected larvae found 

in 13 of 35 habitats.

N 63, 82

 SPECIES

Culex tarsalis Coquillett

(western encephalitis mosquito)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Lagenidium giganteum Couch 

(Pythiales: Pythiaceae)*

1972 UNITED STATES OF 

AMERICA Ex. USA (North 

Carolina). Released in rice 

fields in Colusa County and 

irrigated pastures nr. Hanford, 

California. Recovered 3 

consecutive years but dispersal 

from inoculation sites minimal.

N 44, 91, 152

* Member of the Kingdom Chromista, Phylum Oomycota. All other entries in this list belong to in the Kingdom Fungi.
28 Table C: Exotic fungi and oomycetes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: LEPIDOPTERA

 FAMILY

Lymantriidae

 SPECIES

Lymantria dispar (L.)

(gypsy moth)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Entomophaga maimaiga Humber, 

Shimazu & Soper

 (Zygomycetes: 

Entomophthoraceae)

1910-1911 UNITED STATES OF 

AMERICA Ex. Japan 

(Nishigahara). Released in 

Massachusetts (Boston area). 

Not established.

I 138

1985 UNITED STATES OF 

AMERICA Ex. Japan (Ishikawa 

Prefecture). Released in 

southwestern New York State. 

Not established.

I 60

1986 UNITED STATES OF 

AMERICA Ex. Japan (Ishikawa 

Prefecture). Released 

in northern Virginia. Not 

established.

I 60

1996 BULGARIA Ex. USA 

(Connecticut). Released 

in Levishte (northeastern 

Bulgaria). No infection in 1997.

N 117

1999 BULGARIA Ex. USA 

(Massachusetts). Released 

in Karlovo (central Bulgaria). 

Established but negligible 

control.

N 118

2000 BULGARIA Ex. USA 

(Connecticut). Released in 

Levishte. Infections found in 

2002, 2003 & 2004.

N 117

2002 RUSSIA Ex. USA (Virginia). 

Released in Novosibirsk region. 

Establishment not confirmed.

N 2
Table C: Exotic fungi and oomycetes released and target hosts 29

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SUBCLASS: ACARI

 FAMILY

Eriophyidae

 SPECIES

Eriophyes sheldoni (Ewing)

(citrus bud mite)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Hirsutella thompsonii Fisher 

var. vinacea Samson, McCoy & 

O’Donnell

(Ascomycetes: Anamorph of 

Hypocreales)

1985 ARGENTINA Ex. USA (North 

Carolina). Released on lemon 

trees in Tucuman. Initially 

92% decrease in mites but 

persistence unknown.

I 136, 137

 SPECIES

Eriophyes sheldoni (Ewing), Phyllocoptruta oleivora (Ashmead)

(citrus bud mite, citrus rust mite)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Hirsutella thompsonii Fisher var. 

synnematosa Samson, McCoy & 

O’Donnell

(Ascomycetes: Anamorph of 

Hypocreales)

1985 ARGENTINA Ex. Zimbabwe. 

Released in Tucuman. About 

50% infection for both mite 

species after release but 

persistence unknown. 

I 136, 137

 FAMILY

Tetrancychidae

 SPECIES

Mononychellus tanajoa (Bondar)

(cassava green mite)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Neozygites tanajoae Delalibera, 

Hajek & Humber (prev. referred to 

as Neozygites floridana (Weiser & 

Muma) Remaudiére & Keller)

(Zygomycetes: Neozygitaceae)

1998-1999 BENIN Ex. northeastern 

Brazil. Established, epizootics 

occurring in 2002 & 2003 at 

release sites. Molecular probes 

developed to confirm that 

epizootics were caused by 

exotic strains of the introduced 

pathogen rather than a closely 

related native strain.

I 36, 66
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

30

 ORDER: ORTHOPTERA

 FAMILY

Acrididae

 SPECIES

Various species. Principal targets are Melanoplinae: Dichroplus maculipennis 

(Blanchard), Dichroplus elongatus (Giglio-Tos), Dichroplus pratensis Bruner, 

Scotussa lemniscata Stål

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Paranosema locustae (Canning) 

(= Nosema locustae Canning; = 

Antonospora locustae (Canning))

(Microsporidia: Nosematidae)

1978-82 ARGENTINA Ex. USA (Idaho). 

Released in central Argentina. 

Established and, in 1994-5, 

found 75 km from release 

sites. Epizootics occur with 

accompanying host declines 

but levels of infection in 

susceptible species usually 

average < 10%.

N 84

 ORDER: DIPTERA

 FAMILY

Culicidae

 SPECIES

Culex pipiens quinquefasciatus Say (= C. pipiens fatigans Wiedemann; = C. fatigans Wiedemann)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Pleistophora culicis (= Plistophora 

culicis) Weiser

(Microsporidia: Pleistophoridae)

1967 NAURU Ex. Nigeria (Lagos). 

Establishment not confirmed.

N 82

TABLE D: EXOTIC MICROSPORIDIA RELEASED AND TARGET HOSTS
Table D: Exotic microsporidia released and target hosts 31

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: LEPIDOPTERA

 FAMILY

Pyralidae

 SPECIES

Ostrinia nubilalis (Hübner)

(European corn borer)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Nosema pyrausta (Paillot) (= 

Perezia pyraustae Paillot; = 

Glugea pyraustae (Paillot))

(Microsporidia: Nosematidae)

Unk.- betw. 

1952 & 1960

UNITED STATES OF 

AMERICA Ex. USA (Iowa). 

Infected larvae distributed at 

scattered localities throughout 

Illinois. Disease became 

prevalent and kept host 

populations at low levels.

I 37

 FAMILY

Lymantriidae

 SPECIES

Lymantria dispar (L.)

(gypsy moth)

 BIOLOGICAL CONTROL AGENT

RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Nosema portugal Maddox & 

Vávra (= Microsporidium sp.)

(Microsporidia: Nosematidae)

1986 UNITED STATES OF 

AMERICA Ex. Portugal. 

Released in Maryland. 

Established, low levels of 

infection in 1987 which 

persisted for 3 years.

I 72, 73, 94

1992, 1993 UNITED STATES OF 

AMERICA Ex. Portugal. 

Released in Michigan. Low 

levels of infection during the 

season of release, persistence 

not confirmed.

I 6, 7

Endoreticulatus sp. (= Vavraia

sp.)

(Microsporidia: Pleistophoridae)

1986 UNITED STATES OF 

AMERICA Ex. Portugal. 

Released in Maryland. Not 

established.

I 72, 73, 94
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

32

 ORDER: ORTHOPTERA

 FAMILY

Gryllotalpidae

 SPECIES

Scapteriscus abbreviatus Scudder, Scapteriscus borelli Gigli-Tos, Scapteriscus vicinus Scudder

(mole crickets)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Steinernema scapterisci Nguyen 

& Smart

(Rhabditida: Steinernematidae) 

1985 UNITED STATES OF 

AMERICA Ex. Uruguay. 

Released in Florida. 

Established, host populations 

declined by 85-98%, by 

1988 infected hosts collected 

23 km from release site. 

Establishment on golf courses 

not as successful but > 27% 

reduction in hosts when 

persisting.

I 113

 ORDER: COLEOPTERA

 FAMILY

Scarabaeidae

 SPECIES

Oryctes rhinoceros (L.)

(rhinoceros beetle)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Rhabditis sp.

(Rhabditida: Rhabditidae)

1954 FIJI Ex. Sri Lanka. Results of 

release not reported.

I 28

1957 FIJI Ex. Madagascar. 

Recovered after release, 

persistence not confirmed.

I 145

1957 AMERICAN SAMOA Ex. 

Madagascar. Results from 

release unknown.

I 145

Rhabditis sp. nr. maupasi Seurat 

in Maupas

(Rhabdita: Rhabditidae)

1957 SAMOA (= WESTERN 

SAMOA) Ex. Sri Lanka. Results 

from release unknown.

I 145

TABLE E: EXOTIC NEMATODES RELEASED AND TARGET HOSTS
Table E: Exotic nematodes released and target hosts 33

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1957 AMERICAN SAMOA Ex. Sri 

Lanka. Results from release 

unknown.

I 145

1957 WALLIS ISLAND Ex. Sri Lanka. 

Results from release unknown.

I 145

 FAMILY

Curculionidae

 SPECIES

Sitona discoideus Gyllenhal

(sitona weevil)

BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Heterorhabditis heliothidis (Khan, 

Brooks & Hirschmann)

(Rhabditida: Heterorhabditidae)

1982 AUSTRALIA Ex. New Zealand. 

Released in So. Australia. No 

infections ever found.

I 3

ORDER: DIPTERA

 FAMILY

Culicidae

 SPECIES

Culex pipiens quinquefasciatus Say

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni

Tsai & Grundmann) 

(Mermithidae)

1971-72 TAIWAN Ex. USA (Louisiana). 

Released in Taipei. Infection 

rates low after release and no 

indication of establishment.

N 100

1974 TAIWAN Ex. USA (Louisiana). 

Released in Taipei. Recycling 

occurred through 196 days 

after release but continued 

persistence not confirmed.

N 27

1972 THAILAND Ex. USA 

(Louisiana). Large releases in 

ditches and drains in Bangkok, 

infection 0-27%, no recycling.

N 26 (in 115)

COLEOPTERAScarabaeidaeRhabditis sp. Oryctes rhino. (L.)
34 Table E: Exotic nematodes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Aedes spp., Ochlerotatus spp. (10 species total)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni 

Tsai & Grundmann) 

(Mermithidae)

1974 CANADA Ex. USA (Louisiana). 

Released in snow melt pools 

in Winnipeg, Manitoba. No 

infection.

N 49 (in 115)

1975-76 CANADA Ex. USA (Louisiana). 

Released in snow melt pools in 

Winnipeg, Manitoba. Meager 

parasitism after one winter 

and continued persistence 

questionable.

N 50

 SPECIES

Aedes polynesiensis Marks, Aedes aegypti (L.)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni 

Tsai & Grundmann)

(Mermithidae) 

1978 TOKELAU ISLANDS Ex. 

USA (Louisiana). Released 

on Fakatao Atoll in tree holes 

and man-made containers. 

Established in 35 of 41 

sites with 14-22% infection. 

Persisted at least 3 years.

N/I 83

 SPECIES

Anopheles nyssorhynchus albimanus Wiedemann

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni 

Tsai & Grundmann)

(Mermithidae) 

1983 COLOMBIA Ex. USA 

(Louisiana). Released in El 

Valle. Established and cycled 

over 27 months, effectively 

reduced host population with 

coincident reduction in malaria 

among schoolchildren.

N 127

DIPTERACulicidae
Table E: Exotic nematodes released and target hosts 35

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Anopheline species: Anopheles dthali Patton, Anopheles superpictus Grassi, Anopheles sergentii 

(Theobald), Anopheles turkhudi Liston, Anopheles culicifacies Giles

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni 

Tsai & Grundmann)

(Mermithidae)

1984-85 IRAN Ex. USA (Louisiana). 

Established, 56-61% parasitism 

immediately post-release but 

only minor reductions in host 

populations. 8% parasitism at 1 

of 13 sites 1 year after release. 

Effective long-term control 

unlikely.

N 162

 SPECIES

Anopheles punctipennis (Say), Anopheles crucians Weidemann, Aedes vexans (Meig.), Culex 

restuans Theobald, Culex pipiens L.

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni 

Tsai & Grundmann)

(Mermithidae) 

1975 UNITED STATES OF 

AMERICA Ex. USA (Louisiana). 

Released in Maryland. 

Established, 50-100% host 

mortality even 2 years after 

release.

N 106

 SPECIES

Anopheles freeborni Aitken, Culex tarsalis Coquillett 

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni

Tsai & Grundmann)

(Mermithidae)

1975-76 UNITED STATES OF 

AMERICA Ex. USA (Louisiana).

Released in California rice 

fields. Continuous partial 

control through rice growing 

season with mean weekly 

infection for both species 

> 60%. Survived chemicals, 

drying, harvest, winter and 

cultivation and parasitized 

hosts next summer.

N 115

DIPTERACulicidae
36 Table E: Exotic nematodes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 SPECIES

Anopheles nyssorhynchus albimanus Weidemann, Anopheles punctipennis (Say)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Romanomermis culicivorax Ross 

& Smith (= Reesimermis nielseni

Tsai & Grundmann) 

(Mermithidae)

1977 El SALVADOR Ex. USA 

(Louisiana). Released in 

Lake Apasteque. Releases 

through year yielded 46-96% 

parasitism; up to 17x reduction 

in host populations. Recycling 

not reported, questionable.

N/I? 116, 127

 SPECIES

Culex pipiens quinquefasciatus Say, Aedes aegypti (L.)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Octomyomermis muspratti

Obiamiwe

& Macdonald

(Mermithidae)

1967 NAURU Ex. Zambia. After 

release, parasitism found in 

several tree holes but long-term 

establishment not reported. 

N/I 82, 119

ORDER: LEPIDOPTERA

 FAMILY

Lymantriidae

 SPECIES

Lymantria dispar (L.)

(gypsy moth)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Hexamermis sp.

(Mermithidae)

1974 UNITED STATES OF 

AMERICA Ex. Austria. 

Released in New Jersey. Not 

established.

I 30

1976 UNITED STATES OF 

AMERICA Ex. Japan 

(Hokkaido). Released in 

Pennsylvania. Not established.

I 30

DIPTERACulicidae
Table E: Exotic nematodes released and target hosts 37

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: HYMENOPTERA

 FAMILY

Siricidae

 SPECIES

Sirex noctilio F.

(European woodwasp, Sirex wasp)

 BIOLOGICAL CONTROL AGENT RELEASE 

DATE

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Deladenus siricidicola Bedding (= 

Beddingia siricidicola (Bedding))

(Rhabditida: Neotylenchidae)

1967 NEW ZEALAND Ex. New 

Zealand (North Island). 

Released on South Island. 

29-76% infection reported 

within first year of release and, 

by 1970, natural spread of ca. 

50 km. Releases continued at 

least through 1974. Lack of 

establishment at some sites 

linked with low density host 

populations.

I 170

1970 AUSTRALIA Ex. Hungary. 

Released in Tasmania. 

Established, reached high 

levels of parasitism rapidly. 

In one forest, six years after 

release of 50 parasitized 

females, trees were no longer 

being killed by woodwasps. 

Spread to nearby forests and 

also released in other areas. 

Considered the key biological 

agent controlling Sirex.

I 9, 64

1971 AUSTRALIA Ex. Tasmania 

(plus other locations). 

Released in Victoria. 

Established, dispersed by 

woodwasps in local forests 

and by humans between 

forests. Use of this nematode 

became a cornerstone in 

the National Sirex Control 

strategy. Released over many 

years in many areas; 147,000 

radiata pines inoculated in the 

Green Triangle in 1987 alone. 

With over 20 years of in vitro 

production, strain lost virulence 

resulting in replacement of 

strain used for releases.

I 11, 64
38 Table E: Exotic nematodes released and target hosts

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1987 URUGUAY Ex. New Zealand. 

Established, yielding 18% 

parasitism. 

I 17, 121, 128

1989, 1990, 

1994

BRAZIL Ex. Australia. 

Principally released in 3 

southern provinces. After 

loss of infectivity, new strain 

(Kamona from Tasmania) 

introduced in 1994, yielding 

50-80% parasitism. Overall 

established, parasitism 

levels variable but provided 

substantial control in some 

areas.

I 11, 67, 68

1995 SOUTH AFRICA Ex. Australia. 

Released Kamona strain in 

southwestern Cape Province. 

Established, with 23% 

parasitism reported in 1996. 

In 1998, along with cultural 

control, credited with containing 

the spread of the pest. 

I 150, 151

1999 ARGENTINA Ex. Brazil. 

Released in Patagonia. 

Established, 50-60% parasitism 

reported at release site in 

2000. 

I 76, 77

HYMENOPTERASiricidaeSirex noctilio F.Deladenus siricidicola Bedding
Table F: Accidental introductions of pathogens and nematodes 39

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: LEPIDOPTERA

 FAMILY

Zygaenidae

 SPECIES

Harrisina brillians Barnes & McDunnough

(western grapeleaf skeletonizer)

 BIOLOGICAL CONTROL AGENT

YEAR 

FOUND

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Harrisina brillians Granulovirus 

(HbGV)

(Baculoviridae)

early 1950s UNITED STATES OF 

AMERICA Found in San Diego 

Co., California, probably 

inadvertently introduced with 

parasitoids from Mexico and 

USA (Arizona). Infections 

observed in field and virus 

continually wiped out colonies 

for rearing parasitoids.

I 143, 144

 FAMILY

Pyralidae

 SPECIES

Ostrinia nubilalis (Hübner)

(European corn borer)

 BIOLOGICAL CONTROL AGENT

YEAR 

FOUND

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Nosema pyrausta (Paillot) (= 

Perezia pyraustae Paillot; = 

Glugea pyraustae (Paillot))

(Microsporidia: Nosematidae)

1949 UNITED STATES OF 

AMERICA Probably introduced 

from Europe, possibly with 

parasitoids. First found in New 

Jersey but subsequently found 

throughout the host distribution 

in the US. Occurring commonly, 

epizootics develop with high 

host density and widespread 

spatial distribution of hosts.

I 22, 61, 141

TABLE F: ACCIDENTAL INTRODUCTIONS OF PATHOGENS AND NEMATODES
40 Table F: Accidental introductions of pathogens and nematodes

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 FAMILY

Lymantriidae

 SPECIES

Lymantria dispar (L.)

(gypsy moth)

 BIOLOGICAL CONTROL AGENT

YEAR 

FOUND

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Entomophaga maimaiga Humber, 

Shimazu & Soper

(Zygomycetes: 

Entomophthoraceae)

1989 UNITED STATES OF 

AMERICA Ex. Japan. First 

found in 1989 in 7 northeastern 

states but spread naturally and 

through releases. Established 

in Connecticut, Delaware, 

Maine, Massachusetts, 

Maryland, Michigan, North 

Carolina, New Hampshire, 

New Jersey, New York, Ohio, 

Pennsylvania, Rhode Island, 

Virginia, Vermont, Wisconsin, 

West Virginia and in Ontario, 

Canada. Host populations 

remain low the majority of 

years and sites, although 

localized increases can occur 

infrequently.

I 42, 57, 60, 107

Lymantria dispar Multicapsid 

Nucleopolyhedrovirus (LdMNPV)

(Baculoviridae)

1907 UNITED STATES OF 

AMERICA (probably 

Massachusetts). Thought 

to have been introduced 

after 1900 from Europe with 

parasitoids released for 

classical biological control or 

with plant material and spread 

through the host population. 

Until E. maimaiga became 

established, caused epizootics 

in high density, defoliating 

host populations, resulting in 

spectacular population crashes. 

Spreads naturally after the host 

population spreads into new 

areas.

I 51, 57

LEPIDOPTERA
Table F: Accidental introductions of pathogens and nematodes 41

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 ORDER: HYMENOPTERA

 FAMILY

Siricidae

 SPECIES

Sirex noctilio F.

(European woodwasp, Sirex wasp)

 BIOLOGICAL CONTROL AGENT

YEAR 

FOUND

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Deladenus siricidicola Bedding (= 

Beddingia siricidicola (Bedding))

(Rhabditida: Neotylenchidae) 

1962 NEW ZEALAND (North Island).

Thought to have arrived with 

host. Attributed with being 

the most important agent 

controlling host on the North 

Island. 

I 10, 169

 FAMILY

Diprionidae

 SPECIES

Gilpinia hercyniae (Hartig) (= Diprion hercyniae Hartig)

(European spruce sawfly)

 BIOLOGICAL CONTROL AGENT

YEAR 

FOUND

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Gilpinia hercyniae

Nucleopolyhedrovirus (GhNPV)

(Baculoviridae)

1936 CANADA and UNITED STATES 

OF AMERICA Probably ex. 

Europe. Believed introduced 

with parasitoids. Virus first 

found in New Brunswick and 

then Maine, Vermont and 

New Hampshire, after which 

it spread from south to north 

and first found in Quebec in 

1940. Also transfered to sites in 

Quebec and Ontario but some 

transfers were unsuccessful 

and virus spread on its own. 

By 1942, virus was distributed 

throughout most of the infested 

areas and was credited as 

cause of rapid decline in pest 

outbreak after 1942. Virus plus 

parasitoids appear to have 

permanently solved problems 

due to this pest in eastern 

North America. 

I 5, 21, 34, 35, 

92
42 Table F: Accidental introductions of pathogens and nematodes

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

1970 or 

1971

UNITED KINGDOM (WALES) 

Probably ex. Europe. Spread 

from small epicenter and 

controlled pest outbreak by 

1974. 

I 35, 43

 FAMILY

Formicidae

 SPECIES

Solenopsis invicta Buren

(red imported fire ant)

 BIOLOGICAL CONTROL AGENT

YEAR 

FOUND

COUNTRY, SOURCE, AND 

RESULTS FROM INTRO.

PEST 

ORIGIN REFERENCES

Thelohania solenopsae Knell, 

Allen & Hazard

(Microsporidia: Thelohaniidae)

1996 UNITED STATES OF 

AMERICA Ex. South America. 

Found in Florida. Infects mostly 

polygynous colonies. Chronic 

debilitation of infected queens 

yields smaller colony sizes and 

possibly prolonged death of 

colonies.

I 112, 153

HYMENOPTERADiprionidaeGilpinia hercyniae (Hartig)GhNPV
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

43

Virus

 Family Baculoviridae

 Agrotis segetum Granulovirus (AsGV)

 Anticarsia gemmatalis Multicapsid Nucleopolyhedrovirus (AgMNPV)

 Gilpinia hercyniae Nucleopolyhedrovirus (GhNPV)

 Harrisina brillians Granulovirus (HbGV)

 Lymantria dispar Multicapsid Nucleopolyhedrovirus (LdMNPV)

 Lymantria monacha Nucleopolyhedrovirus

 Neodiprion sertifer Nucleopolyhedrovirus (NeseNPV)

 Pseudoplusia includens Singlecapsid Nucleopolyhedrovirus (PiSNPV)

 Trichoplusia ni Nucleopolyhedrovirus (TnNPV)

 Family Unassigned

 Oryctes rhinoceros Virus (Orv)

Domain Bacteria

 Phylum Firmicutes

 Class Bacilli

 Order Bacillales

 Family Bacillaceae

 Paenibacillus popilliae

Domain Eukarya

 Kingdom Fungi

 Phylum Chytridiomycota

 Class Chytridiomycetes

 Order Blastocladiales

 Family Coelomomycetaceae

 Coelomomyces stegomyiae

 Phylum Zygomycota

 Class Zygomycetes

 Order Entomophthorales

 Family Entomophthoraceae

 Entomophaga grylli 

 Entomophaga maimaiga

 Pandora neoaphidis

 Zoophthora radicans

 Family Neozygitaceae

 Neozygites fresenii

Neozygites parvispora

 Neozygites tanajoae

CHART 1: CLASSIFICATION OF PATHOGENS AND NEMATODES PURPOSEFULLY 

INTRODUCED FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES OR 

ESTABLISHED AFTER ACCIDENTAL INTRODUCTION 1
44 Chart 1: Classification of pathogens and nematodes

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 Phylum Ascomycota

 Class Loculoascomycetes

 Order Pleosporales

 Family Tubeufiaceae

 Podonectria coccophila

 Class Discomycetes

 Order Rhytismatales

 Family Triblidiaceae

 Triblidium caespitosum 

 Class Pyrenomycetes

 Order Hypocreales (anamorphic/asexual forms)

 Aschersonia aleyrodis

Aschersonia goldiana

 Aschersonia spp.

 Beauveria bassiana

 Beauveria brongniartii

 Fusarium coccophilum

 Fusarium juruanum

 Hirsutella thompsonii var. synnematosa

 Hirsutella thompsonii var. vinacea

 Lecanicillium lecanii

 Metarhizium anisopliae

 Phylum Microsporidia

 Class Microsporea

 Order Nosematidida

 Family Nosematidae

 Nosema portugal

 Nosema pyrausta

 Paranosema locustae

 Order Microsporida

 Family Pleistophoridae

 Endoreticulatus sp.

Pleistophora culicis

 Family Thelohaniidae

 Thelohania solenopsae

 

 Kingdom Chromista (= Kingdom Straminipila)

 Phylum Oomycota

 Class Oomycetes

 Order Pythiales

 Family Pythiaceae

 Lagenidium giganteum
Chart 1: Classification of pathogens and nematodes 45

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 Kingdom Animalia

 Phylum Nematoda

 Class Chromadorea

 Order Rhabditida

 Family Steinernematidae

 Steinernema scapterisci

 Family Neotylenchidae

 Deladenus siricidicola

 Family Rhabditidae

 Rhabditis sp.

Rhabditis sp. nr. maupasi

 Family Heterorhabditidae

 Heterorhabditis heliothidis

 Class Enoplea

 Order Mermithida

 Family Mermithidae

 Hexamermis sp.

Octomyomermis muspratti

Romanomermis culicivorax

1

 Organization of domains based on Woese et al. (156). Placement of the Phylum Microsporidia in the Kingdom 

Fungi is a recent change (75) and, while we have adopted this change here, the higher order affiliation of this group 

is still somewhat in a state of flux.Nematode classification based on De Ley & Blaxter (38).
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

46

Kingdom Animalia

 Phylum Arthropoda

 Class Insecta

 Order Orthoptera

Family Gryllotalpidae

 Scapteriscus abbreviatus

 Scapteriscus borelli

 Scapteriscus vicinus

 Family Acrididae

 Camnula pellucida

 Dichroplus elongatus

 Dichroplus maculipennis

 Dichroplus pratensis

 Melanoplus bivittatus

 Melanoplus sanguinipes

 Phaulacridium vittatum

 Scotussa lemniscata

 Order Hemiptera

 Family Cercopidae

 Aeneolamia flavilatera

Family Cicadellidae

 Empoasca fabae

 Family Aphididae

 Aphis gossypii

 Macrosiphum solanifolii

 Metopolophium dirhodum

 Therioaphis maculata

 Family Aleyrodidae

 Aleurodicus cocois

 Aleurothrixus floccosus

 Dialeurodes sp.

Dialeurodes citri

Singhiella citrifolii

 Family Coccidae

 Ceroplastes rubens

Coccus viridis

 Eucalymnatus tessellatus

 Family Diaspididae

 Aonidiella aurantii

 Aspidiotus destructor

 Cornuaspis beckii

 Quadrispidiotus perniciosus

 Order Thysanoptera

 Family Thripidae

 Thrips tabaci

CHART 2: CLASSIFICATION OF INSECTS AND MITES TARGETED BY CLASSICAL BIOLOGICAL 

CONTROL PROGRAMS USING PATHOGENS OR NEMATODES, OR HOSTS OF ACCIDENTAL 

INTRODUCTIONS OF PATHOGENS OR NEMATODES
Chart 2: Classification of insects and mites 47

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 Order Coleoptera 

 Family Scarabaeidae

 Adoretus tenuimaculatus

 Alissonotum impressicolle

 Anoplognathus sp.

 Cochliotis melolonthoides

Dermolepida albohirtum

 Lepidiota sp.

 Lepidiota pruinosa

 Leucopholis irrorata

 Oryctes monoceros

Oryctes rhinoceros

 Papuana huebneri

 Phyllophaga smithi

 Scapanes australis

 Schizonycha sp.

 Family Curculionidae

 Otiorhyncus arcticus

Otiorhyncus nodosus

 Sitona discoideus 

Order Diptera

 Family Culicidae

 Aedes spp.

Aedes aegypti

Aedes polynesiensis

 Aedes vexans

 Anopheles crucians

 Anopheles culicifacies

 Anopheles dthali

 Anopheles freeborni

 Anopheles nyssorhynchus albimanus

 Anopheles punctipennis

 Anopheles sergentii

 Anopheles superpictus

 Anopheles turkhudi

 Culex pipiens

 Culex pipiens quinquefasciatus

 Culex restuans

 Culex tarsalis

 Ochlerotatus spp.

Order Lepidoptera

Family Zygaenidae

 Harrisina brillians

 Family Pyralidae

 Ostrinia nubilalis

Family Lymantriidae

 Lymantria dispar

 Lymantria monacha
48 Chart 2: Classification of insects and mites

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

 Family Noctuidae

 Agrotis segetum

Anticarsia gemmatalis

 Pseudoplusia includens

 Trichoplusia ni

 Order Hymenoptera

 Family Siricidae

 Sirex noctilio

 Family Diprionidae

 Gilpinia hercyniae

Neodiprion sertifer

 Family Formicidae

 Solenopsis invicta

 Class Arachnida

 Subclass Acari

 Order Prostigmata

 Family Eriophyidae

 Eriophyes sheldoni

 Phyllocoptruta oleivora

 Family Tetranychidae

 Mononychellus tanajoa
CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

49

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CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

56

Dichroplus pratensis 30

Diprion hercyniae 10, 41-42

E

Empoasca fabae 14

Eriophyes sheldoni 29

Eucalymnatus tessellatus 20

G

Gilpinia hercyniae 10, 41-42

H

Harrisina brillians 6, 39

L

Lachnosterna smithi 24

Lepidiota pruinosa 25

Lepidiota sp. 25

Lepidosaphes beckii 21

Leucopholis irrorata 25

Lymantria dispar 6, 28, 31, 36, 40

Lymantria monacha 7

M

Macrosiphum solanifolii 17

Melanoplus bivittatus 13

Melanoplus sanguinipes 13-14

Metopolophium dirhodum 16

Mononychellus tanajoa 29

N

Neodiprion sertifer 9-10

O

Ochlerotatus spp. 34

Oryctes monoceros 5

Oryctes rhinoceros 3-5, 11, 23, 32-33

Ostrinia nubilalis 31, 39

Otiorhynchus arcticus 26

Otiorhynchus nodosus 26

P

Papuana huebneri 11, 24

Phaulacridium vittatum 13

Phyllocoptruta oleivora 29

Phyllophaga smithi 24

Phytalus smithi 24

Pseudoplusia includens 8

A

Adoretus tenuimaculatus 26

Aedes aegypti 34, 36

Aedes polynesiensis 27, 34

Aedes spp. 34

Aedes vexans 35

Aeneolamia flavilatera 14

Agrotis segetum 8

Aleurodicus cocois 19

Aleurothrixus floccosus 19

Alissonotum impressicolle 24

Anopheles crucians 35

Anopheles culicifacies 35

Anopheles dthali 35

Anopheles freeborni 35

Anopheles nyssorhynchus albimanus 34, 36

Anopheles punctipennis 35-36

Anopheles sergentii 35

Anopheles superpictus 35

Anopheles turkhudi 35

Anoplognathus sp. 25

Anticarsia gemmatalis 7

Aonidiella aurantii 21

Aphis gossypii 16

Aspidiotus destructor 22

Aspidiotus perniciosus 22

C

Camnula pellucida 13

Ceroplastes rubens 20

Clemora smithi 24

Coccus viridis 19-20

Cochliotis melolonthoides 12

Cornuaspis beckii 21

Culex fatigans 30

Culex pipiens 35

Culex pipiens fatigans 30

Culex pipiens quinquefasciatus 30, 33, 36

Culex restuans 35

Culex tarsalis 27, 35

D

Dermolepida albohirtum 25

Dialeurodes citri 18

Dialeurodes citrifolii 17

Dialeurodes sp. 18

Dichroplus elongatus 30

Dichroplus maculipennis 30

Scientific names of insect and mite 

hosts

Index
Index 57

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

Q

Quadraspidiotus perniciosus 22

S

Scapanes australis 6

Scapteriscus abbreviatus 32

Scapteriscus borelli 32

Scapteriscus vicinus 32

Schizonycha sp. 11

Scotussa lemniscata 30

Singhiella citrifolii 17

Sirex noctilio F. 37-38, 41

Sitona discoideus 26, 33

Solenopsis invicta 42

T

Therioaphis maculata 15

Thrips tabaci 23

Trichoplusia ni 8

Scientific names of pathogens and 

nematodes

A

Agrotis segetum Granulovirus (AsGV) 8

Anticarsia gemmatalis Multicapsid Nucleopolyhedrovirus (AgMNPV) 7

Antonospora locustae 30

Aschersonia aleyrodis 19

Aschersonia goldiana 17

Aschersonia spp. 18

B

Baculovirus oryctes 3-6

Beauveria bassiana 24, 26

Beauveria brongniartii 25

Beauveria densa 24

Beddingia siricidicola 37-38, 41

Botrytis tenella 25

C

Cephalosporium lecanii 24

Coelomomyces stegomyiae 27

D

Deladenus siricidicola 37-38, 41

E

Endoreticulatus sp. 31

Entomophaga grylli 13-14

Entomophaga maimaiga 28, 40

F

Fusarium coccophilum 21-22

Fusarium episphaerea 21

Fusarium juruanum 22

G

Gilpinia hercyniae Nucleopolyhedrovirus (GhNPV) 

10, 41-42

Glugea pyraustae 31, 39

H

Harrisina brillians Granulovirus (HbGV) 6, 39

Heterorhabditis heliothidis 33

Hexamermis sp. 36

Hirsutella thompsonii 29

Families of insect and mite hosts

Acrididae 13-14, 30

Aleyrodidae 17-19

Aphididae 15-17

Cercopidae 14

Cicadellidae 14-15

Coccidae 19-20

Culicidae 27, 30, 33-36

Curculionidae 26, 33

Diaspididae 21-22

Diprionidae 9-10, 41-42

Eriophyidae 29

Formicidae 42

Gryllotalpidae 32

Lymantriidae 6-7, 28, 31, 36, 40

Noctuidae 7-8

Pyralidae 31, 39

Scarabaeidae 3-6, 11-12, 23-26, 32-33

Siricidae 37-38, 41

Tetranychidae 29

Thripidae 23

Zygaenidae 6, 39
58 Index

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

L

Lagenidium giganteum 27

Lecanicillium sp. 17, 19

Lecanicillium lecanii 20

Lymantria dispar Multicapsid Nucleopolyhedrovirus 

(LdMNPV) 6, 40

Lymantria monacha Nucleopolyhedrovirus 7

M

Metarhizium anisopliae 14, 23-26

Microsporidium sp. 31

Myrangium duriaei 21

N

Nectria flammea 21-22

Neodiprion sertifer Nucleopolyhedrovirus (NeseNPV) 

9-10

Neozygites floridana 29

Neozygites fresenii 16

Neozygites parvispora 23

Neozygites tanajoae 29

Nosema locustae 30

Nosema portugal 31

Nosema pyrausta 31, 39

O

Octomyomermis muspratti 36

Oryctes rhinoceros virus (OrV) 3-6

P

Paenibacillus popilliae 11-12

Pandora neoaphidis 16

Paranosema locustae 30

Perezia pyraustae 31, 39

Pleistophora culicis 30

Plistophora culicis 30

Podonectria coccicola 21

Pseudomicrocera henningsii 22

Pseudoplusia includens Singlecapsid Nucleopolyhedrovirus (PiSNPV) 8

R

Reesimermis nielseni 33-36

Rhabdionvirus oryctes 3-6

Rhabditis sp. 32

Rhabditis sp. nr. maupasi 32-33

Romanomermis culicivorax 33-36

Higher classification of pathogens and 

nematodes

Bacillaceae 11-12

Baculoviridae 6-10, 39, 40-42

Coelomomycetaceae 27

Entomophthoraceae 13-16, 28, 40

Heterorhabditidae 33

Hypocreales 14, 17-26, 29

Mermithidae 33-36

Neotylenchidae 37-38, 41

Neozygitaceae 16, 23, 29

Nosematidae 30-31, 39

Pleistophoridae 30-31

Pythiaceae 27

Rhabditidae 32-33

Steinernematidae 32

Thelohaniidae 42

Triblidiaceae 21

Tubeufiaceae 21

Unassigned virus family 3-6

S

Sphaerostilbe coccophila 21-22

Steinernema scapterisci 32

T

Thelohania solenopsae 42

Triblidium caespitosum 21

Trichoplusia ni Nucleopolyhedrovirus (TnNPV) 8

V

Vavraia sp. 31

Verticillium sp. 17, 19

Verticillium lecanii 20

Z

Zoophthora radicans 14-15
Index 59

CATALOGUE OF INTRODUCTIONS OF PATHOGENS AND NEMATODES FOR CLASSICAL BIOLOGICAL CONTROL OF INSECTS AND MITES

Source countries and regions

Argentina 23

Australia 13-15, 25, 38

Austria 8, 36

Brazil 7, 14, 16, 29, 38

Canada 9-10

China 18

Cuba 18

Europe 39-42

Faroe Islands 26

Fiji 6, 15

France 25-26

Germany 7

Guatemala 8

Hungary 37

India 4-5, 18, 20

Israel 15

Japan 18, 28, 36, 40

Java 23

Madagascar 32

Malaysia 3, 5

Mexico 6, 39

New Zealand 33, 37-38

Nigeria 30

Papua New Guinea 11, 24

Philippines 5

Portugal 31

Praslin Island group 5

Samoa 3-5, 23, 25

Serbia 6

Sierra Leone 22

Singapore 27

Solomon Islands 11

South America 42

Sri Lanka 20, 32-33

Sweden 7, 9

Switzerland 23

Tanzania 5

Tasmania 37

Trinidad 14, 18

Unknown 19-20, 24, 26

Uruguay 32

USA 6, 8, 10-13, 16-19, 21-22, 24, 27-31, 33-36, 39

Vietnam 18

Zambia 36

Zimbabwe 29

Release countries and regions 

(capitalized as in the text)

AMERICAN SAMOA 4, 11, 32-33

ARGENTINA 21, 29-30, 38

AUSTRALIA 13, 15, 25-26, 33, 37

AZERBAIJAN 18

BARBADOS 23

BELGIUM 16

BENIN 29

BERMUDA 17-18, 21

BRAZIL 38

BULGARIA 28

CANADA 9-10, 34, 41

COLOMBIA 8, 34

DENMARK 7-8

El SALVADOR 36

FIJI 3, 26, 32

GEORGIA 18

GUYANA 14

ICELAND 26

INDIA 4-5

IRAN 35

KENYA 11

KIRIBATI 11, 24

MALDIVES 5

MAURITIUS 4, 24

NAURU 30, 36

NEW ZEALAND 37, 41

PALAU 3, 11

PAPUA NEW GUINEA 4

PHILIPPINES 25

RUSSIA 28

SAMOA 3, 23, 32

SARDINIA 6

SEYCHELLES 5, 20, 22

SOLOMON ISLANDS 6

SOUTH AFRICA 38

TAIWAN 24, 33

TANZANIA 5, 11

THAILAND 33

TOKELAU ISLANDS 3, 23, 27, 34

TONGA 4, 23

UNITED KINGDOM (SCOTLAND) 10

UNITED KINGDOM (WALES) 42

UNITED STATES OF AMERICA 6-10, 13-17, 19-22, 

27-28, 31-32, 35-36, 39-42

URUGUAY 38

VIRGIN ISLANDS 19

WALLIS ISLAND 4, 23, 33