Ylo. AiU

Parasites and Predators

of Forest Insects

Liberated in the United States

Through 1960

LIBRARY

RECEIVED

JUL3 0 1962

1 i ft^tm « mmm

BEIT;'ALLE BRANCH

Agriculture Handbook No. 226

U.S. Department of Agriculture

Forest Service

July 1962
Parasites and Predators

of Forest Insects

Liberated in the United States

Through 1960

by Philip B. Dowden

Agriculture Handbook No. 226 J'^y ^'^^

U. S. Department of Agriculture • Forest Service

F«, .ale by the S»p.ri»U,nde». of Docum»..,, V.S. G. »e». PrioU»8 Office

Washington 25, D.C. - Price 30 cent.
CONTENTS

Page

A record of liberations listed under host species 2

Asterolecanium bambusae (Bdv.) and Asterokcanium variolosum (Katz.) 2

Ohermes piceae Eatz 2

Ghoristoneura fumiferana (Clem.) 3

Cnidocampa flavescens (Wlkr.) 3

Coleophora laricella (Hbn.) 9

Dendroctontis Jrontalis Zimm 9

Diprion frutelorum (F.) 9

Diprion hercyniae (Htg.) 9

Diprion similis (Htg.) 16

Oalerucella xanthomelaeTM (Sehr.) 16

Oossyparia spuria (Mod.) 17

Hemerocampa leucostigma (J. E. Smith) 17

Hemeroeampa pseudotsugata McD 17

Hemileuca oliviae CUl 20

Heterarthrus nemoratus (FaU.) 20

Malacosoma disstria Hbn 20

Malacosoma Jragilis (Stretch) 20

Miscellaneous releases of Calosoma 23

Miscellaneous releases of Compsilura 23

Neodiprion abietis (Harr.) 23

Neodiprion fulviceps (Cress.) 23

Neodiprion lecontei (Fitch) 23

Neodiprion pratti pratti (Dyar) 26

Neodiprion pratti paradoxicus Ross 26

Neodiprion sertijer (Geoff.) 26

Neodiprion swainei MXaa. 27

Nygmia phaeorrhoea (Donov.) 27

Porthetria dispar (L.) 27

Pristiphora eriehsonii (Htg.) ^2

Bhyacionia buoliana (Schm.) ^^

Bkyacionia frustraría bushneüi (Busck) 37

Setora (Miresa) nitens Wlk "¿L

Stilpnotia salicis (L.)
Page

Individual species successfully established in the United States. 47

Agathis pumilus (Ratz.) (Braconidae) 47

Anastatus disparis Ruschka (Eupelmidae) 47

ApaTiieles lacteicolor Vier. (Braconidae) 48

Apúnteles melanoscelus (Ratz.) (Braconidae) 48

Apanteks solitarius (Ratz.) (Braconidae) 49

Aphideeta obliterata (L.) (CoccineUidae) 49

Aphidoletes thompsoni Mohn (Cecidomyiidae) 49

Aptesis basizonia (Grav.) (Ichneumonidae) 50

Blepharipoda scutellata Qi.-D.) (Tachinidae) SO

Calosoma sycophanta (L.) (Carabidae) 51

Campoplex frustranae Cush. (Ichnevunonidae) 51

Carabus spp. (Carabidae) 52

Oarcelia laxifrons Yñl. (Tachinidae) 52

Ohaetexorista javana a. & B. (Tachinidae) 52

Chrysocharislaricinellae (Ratz.) (Eulophidae) 53

Coccophagus insidialor CDalm.) (Eidophidae) 53

Compsilura concinnata (Meig.) (Tachínidae) 54

Oopidosoma geniculatum Dalm. (Encyrtidae) 54

Gremifania nigrocellviata Cz. (Chamaemyiidae) 54

Dahlbominus Juscipennis (Zett.) (Eulopludae) 55

Drino (Prostnrmia) bohémica Mesn. (Tachinidae) 56

Erynniopsis rortdani Towns. (Tachinidae) 56

Eupteromalus nidulans (Thomson) (Pteromalidae) 57

Exenterus abruptorius (Thnb.) (Ichneumonidae) 57

ExeTiierus spp. (Ichneumonidae) 57

Exorista larvamm (L.) (Tachinidae) 58

Laricobius erichsoni Roseuh. (Derodontidae) 58

Leucopia {Neoleucopis) obscura (Hal.) (Chamaemyiidae).. 59

Mesoieius tenthredinis M-orley (Ichneumonidae) 60

Meteorus versicolor (Wesm.) (Braconidae) 60

Monodontomerus aereus Walk. (TorjTnidae) 61

Monodontomerus dentipes (Dalman) (Torymidae) 61

Ooencyrtus kuwanai (How.) (Encyrtidae) 61

Orgilus obscurator (Nees) (Ichneumonidae) 62

Parasetigena silvestrisH.-'D. (Tachinidae) 62

Phanomeris phyllotomae Mues. (Braconidae) 63

Phobocampe disparis (Vier.) (Ichneumonidae) 63

Phytodietus fwni^erarme Roh. (Ichneumonidae) 64

Pimpla turioneliae (L.) (Ichneumonidae) 64

Scymnus (Pullus) impexus (Muls.) (CoccineUidae) 64

Temelucha interruptor {GT&V.) (Ichneumonidae) 65

Tetrastichus brevistigma Gahan (Eulophidae) 65

Tetrastiekus turionum (Htg.) (Eulopludae) 65

TownsendieUomyia nidicola (Tns.) (Tachinidae) 66

Index of host species 67

Index of insect enemies 69
Parasites and Predators of Forest Insects

Liberated in the United States

Through 1960

by Philip B. Dowden '

For over 60 years parasites and predators of forest insect pests

have been imported and released in the United States. Most of the

records of this work are concentrated at the Northeastern Forest

Experiment Station's Forest Insect Laboratory, New Haven, Conn.

With time these files become more and more obscure, and similar

data are being filed elsewhere. Because this information is often

important, the purpose of this handbook is to summarize all work

of this nature in the United States through 1960.

Canada has done more in introducing and liberating parasites and

predators of forest insects than has the United States; therefore,

Canadian reports should be referred to for a complete record of work

done against pests occurring in both countries. The Canadian work

is briefly mentioned in this publication under the principal host

species.

The subject matter is presented in two sections, in much the same

way that it was in the report by C. P. Clausen, Biolorical Control of

Insect Pests in the Continental United States (U.S.D.A. Technical

Bulletin 1139, June 1956), which included all importations and establishments in this country through 1950. Data for forest and shade

tree insects only are included in this handbook, thus presentmg m

greater detail the work abeady done with each species.

In the first section, all species that have been imported and hberated are listed under their hosts, which are arranged alphabetically.

In the second section, a short sketch is presented about each species

successfully established. A few species are included that have been

recovered in Canada but not in the United States, and there may

be others successfully established but not yet recovered. Untortunately, attempts to recover introduced species have varied greatly.

For some species almost no attempt has been made for recovery, and

in others it is stiU too early to determine whether a species is successfuly estabUshed or not. No evaluation is made of the eflectiveness

of the numerous biotic factors that operate against a single Host.

However, figures regarding "percentage parasitization are irequently ilsedT for they often tell much about the relative unportance

of individual species in the work of control.

• Entomologist, Nortieastem Forest Eipcrimmt Station Forest Service^ íf¿-g^S'sërvta!''Brtb Insect Mentlñoation and Parasite Introduction Kescaroh Brancli, AETiculturu Besearcn oervu»,.

agencies are units of the U.S. Department of Agriculture.
2 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

Nomenclatural and taxonomic difficulties and divergent views in

the dipterous family Tachinidae have made it seem best to record

most of the species under the names or combinations used in 1956

when Technical Bulletin 1139 was published. Where these names

differ from those now commonly used, changes are indicated in the

text.

The general plan of listing a parasite or predator according to the

host species against which it was released has been adhered to, but

it should be kept in mind that many parasites and predators attack

a number of host species. The index cites those species liberated

against more than one host.

In the tables, an asterisk shows that a species is known to be

established. Under "Origin" the source of the first specimens received

and liberated is given. Under "Living Insects Released" the numbers

of males and females are shown separately where records by sex are

available.

References to literature have been omitted except where noted

in the text.

A RECORD OF LIBERATIONS LISTED

UNDER HOST SPECIES

Asterolecanium hambusae (Bdv.), bamboo scale, and

Asterolecanium variolosum (RatzOs golden oak scale

Small collections of Asterolecanium variolosum (Ratz.) were made

in New England and sent to Puerto Rico in 1937 and to California

Ln 1952. The scales were parasitized by Habrolepis dalmani (West.),

and it was planned to release the adult parasites, when they emerged,

against the bamboo scale in Puerto Rico and the golden oak scale in

California. So far as is known the parasite did not become established

in either area.

Chermes piceae Ratz. =Adelges piceae (Ratz.)

balsam woolly aphid

Canadian entomologists introduced several predators of the balsam

woolly aphid, Chermes piceae Ratz., from Europe between 1933 and

1937 for release in the Maritime Provinces. Leucopis (Neoleucopis)

obscura (Hal.) was the only one that became establiáied at that time.

The work was reactivated on a much larger scale in Europe in 1951.

Since 1957, collectors have also searched for predators in Australia,

Pakistan, India, and Japan. The work in Japan was done by the

U.S. Forest Service in the summer of 1958.

Several European predators in addition to Leucopis have now been

successfully established in Canada. Recently, with the discovery

that the balsam woolly aphid is present in Washington and Oregon

and also in North Carolina, colonies of imported material have been

made available for release in these new infestations. In 1959 a number
PARASITES AND PREDATORS OF FOREST INSECTS 3

of species, which were introduced for the first time from Pakistan,

were released in the Pacific Northwest.

At present four predators are Imown to be established in the

Canadian Maritimes. They are LeiKopis obscura (Hal.), Larwohius

eñehsoni Rosenh., Scymnus (Pidlus) imperas (Muís.), and Oremifania

nigrocellidata Cz. The first two are also established in New England.

Leucopis spread naturally into Maine; and from there it has been

recolonized at several points in New England and the Pacific Northwest. Laricobius was recovered at a liberation point in southern

Vermont, and it is anticipated that it wiU be recovered in Maine

from large liberations made in the summer of 1959.

Leucopis, Laricobius, and a Cecidomyiid (Aphidoletes thompsoni

Mohn) have been established in the Pacific Northwest. A. thompsoni

has not yet been established in Canada.

The predators that have been liberated against the balsam woolly

aphid are recorded in table 1.

Chorístoneura fumiferana (Clem.)

spruce budworm

Prom studies made during the outbreak of the spruce budworm,

Choristoneura Jumiferana (Clem.), of 1913-22 in eastern Canada, it

was learned that several western budworm parasites did not occur in

the East. Therefore, Canadian workers reared larvae collected in

British Columbia to obtain these parasites for release ia eastern

infestations from 1943-47. The United States conducted similar

work in Colorado and Oregon from 1945-50. The two countries

cooperated in some of this work (table 2). One western species,

Phytodietus fumijeranae Roh., was propagated for several seasons in

both Canada and the United States, particularly by the State of

Maine, to obtain colonies for liberation. Oddly enough, none of the

western parasites seem to have become established in the East.

Canada also imported from Europe parasites of a closely allied

species, Choristoneura muriana (Hbn.). A number of these parasites

were released in Canada, but none seem to have become established.

Cnidocampaflavescens (Wllir.)

oriental moth

In 1929 and 1930, large shipments of cocoons of the oriental moth,

Onidocampa flavescens (Wlkr.), heavUy parasitized by Chaetexonsta

javana B. & B., were sent to the Gypsy Moth Laboratory at Meh-ose

Highlands, Mass., from Japan. The parasite was established a,ncl

has become very effective throughout the known axea of mtestation

in the environs of Boston, Mass. (table 3). A second parasite, Chrysu

shanghaiensis Smith, which was released mWVJ, was recovered m

1919 in small numbers, but it apparenüy failed to persist for it has

not been recovered since then.
TABLE 1.—^Liberations against the balsam woolly aphid, Chermes piceae Eatz.

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

d'à" 9 Î Total

India __.

---do

Europe

do

Pakistan

JapanEurope

Pakistan

do

Portland, Oreg

Monmouth, Oreg

Portland, Oreg

Santiam Pass, Oreg

Mount Mitchell, N.C..

Corvallis, Oreg

Green River, Wash

Wind River, Wash

Bradley, Maine

Asheville, N.C

Black Rock, Oreg

Corvallis, Oreg

McKenzie Pass, Oreg__

Monroe, Greg _

1960

1959

1959

1959

1960

1958

1958

1959

1959

1959

1957, 1959

1957

1959

1968, 1969

1959

1957

1967, 1969

1959

1959

1958

1958

1959

1969

1968

1960

1959

1960

1960

1959

1960

1960

India

do

do

do

Europe

IlllldClIIIIII

do

do

do.

do

do

do -

do

do

do

... do

26

17

66

21

1,300

396

326

718

"2Í,'2Í6'

26

14

var. bispilota Wse. 54

21

1,230

396

325

718

*Avhidoletes thompsoni

Mohn ""3,"236"

4,357

68

1,348

1,149

3,201

124

1,576

742

""5;"573"

7,163

132

2,502

1,485

5,961

244

1,984

1,320

1,185

8,809

11, 620

200

3,850

2 634

Willamette Pass, Oreg—

Green River, Wash...

Coldwater Creek,

Wash.

Handle, Wash

9,162

368

3, 560

do

Pakistan

Japan

do

Europe

IlllldoIIIIIII

Pakistan.

do

do

do

do

do

do

85"

55

80

159

830

385

44

316

300

2,400

1,641

1, 100

925

2 062

Ballia euckaris Muls Black Rock, Oreg

Monroe, Oreg

86

Chilocorus kuwanae Silv.- 55

Coldwater Creek,

Wash.

Corvallis, Oreg

80

Cremifania nigrocellulata 159

Cz. 830

Willamette Pass, Oreg..

Santiam Pass, Oreg

Corvallis, Oreg

Portland, Oreg

Santiam Pass, Oreg

Green River, Wash

Toutle River, WashV...

385

Exochomus Uturatus 44

Corh.

Exochomus uropygialis 316

Muls. 300

2 400

1,641

1,100

925
India

Europe

India

Pakistan

Europe

do

Australia

India

Portland, Oreg_

Bradley, Maine

Mount Mitchell, N.C..

do

1959

1959

1959

1960

1965

1968

1958

1959

1958

1955

1958

1958

1958

1959

1959

1959

1960

1937

1955

1956

1969

1954

1956

1956

1958

1960

1960

1955

1960

1960

1960

1959

1959

1959

India

Europe

IlllldoIIIIIII

do

do

do

do

do

do

do

do

do

India

do

Pakistan

New England.

New Brunswick

New England.

do

do-

do

do

do

do.

Europe

IlllldoIIIIIII

Australia

do

do

do

India

do

11

15, 968

624

1,200

500

1,675

1,705

1,640

1,200

500

1,600

1,320

1,150

169

200

142

569

888

622

2,085

452

730

620

700

300

400

270

2,000

1,400

2,000

859

57

91

10

16, 210

619

1,100

494

s Black Rock, Oreg

Corvallis, Oreg

1,56(

1, 70S

1,600 ' ? Willamette Pass. Oreg.. 1,200

489

Green River, Wash

Coldwater Creek,

Wash.

Wind River, Wash

Monmouth, Oreg

Willamette Pass, Oreg..

1,600

1,320

1,150

Leu dimidiata (F.) 15£

Leucopií {Neoleucopis)

n.sp.

*Leucopis obscura (Hal.).

15

Mount MitcheU, N.C..

Grantham, N.H

Lincoln, N.H

142

£69

888

Cedar River, N.Y

Willamette Pass, Oreg..

485

2,085

462

Texas FaUs, Vt

WaUingford, Vt

Green River, Wash

Mount Mitchell, N.C..

Monroe, Oreg

730

620

700

Seymnus (.Pullua)

impexue (Muís.)

290

400

Somerset, Vt 268

Scymnua {Diomus)

jmmüio Wse.

Mount Mitchell, N.C..

do

Green River, Wash

Toutle River, Wash

Santiam Pass, Oreg

Rändle, Wash

1,900

1, 400

2,000

859

Synharmonia conglóbala 30

(L.) 91

*Kiiown to be established.
TABLE 2.—Liberations against the spruce budworm, Ghonstoneura jumiferana (Clem•)

Num. Livinff insects released

Species Origin Where liberated Year Where

collected

ber

shipped

cf c? 9 9 Total

Agria affinis (Fall.) ' British

Columbia.

Castle Hill, Maine 1948 British

Columbia.

335 154 169 323

Harrietstown, N.Y 1947 do 500 247 247 494

North Elba, N.Y 1947 do 500 245 250 495

Santa Clara, N.Y. .. 1946 do 50 20 24 44

Ceromasia auricaudata united States. Castle HUl, Maine

New Canada, Maine

1948

1950

Oregon

do

46

122

46

Tns. 70 52 122

T5 R16, Maine _ 1947 Colorado 1,232 593 625 1,218

T16 R5, Maine 1949 Oregon 959 467 492 959

Altamont, N.Y 1946 Colorado 483 150 316 466

Brighton, N.Y 1946 do 962 464 498 962

Colton, N.Y... 1946 do 1 020 520 500 1 020

Harrietstown, N.Y 1947 do '221 113 108 221

do 1947 British

Columbia.

500 249 247 496

Long Lake, N.Y 1946 Colorado 951 451 500 951

North Elba, N.Y 1946 do 1,000 600 500 1,000

Santa Clara, N.Y 1946 do 700 200 500 700

Wilmington, N.Y 1946 do 500 250 250 500

Compsilura eoncinnata

(Meig.)

Europe Unoompaghre N. F.,

Colo.

1947 New England. ' 1, 000 70 70

El Rito, N. Mex 1947 do 434 78 353 431

United States.

Taos, N. Mex 1947

1947

do.

Colorado

430

30

160 lioplectis > 4-cingulatxis

TProv.).

Colton, N.Y 30

Phytodietus fumiferanae British Castle HUl, Maine 1948 British 200 50 150 200 Columbia. Columbia.

Hamlin, Maine 1964 Lab. reared 400 50 350 400 New Canada, Maine 1951 do 80 27 56 83 do 1952 do 164 66 98 164
Portage, Maine

Stockholm, Maine

19SS

1955

do

do

3S

830

35

420 410 830

Til E7, Maine 1955

1954

do

do

39

295

37

T13R12, Maine 295 295

T14R6, Maine 1956 do 70 70 70

T14R6, Maine 1957 do 30 SO 30

T14 R9, Maine 1953 do 67 63 63

T16 R4, Maine 1951 do 152 30 122 152

T16R4, Maine 1952 do 212 110 102 212

T16 R5, Maine 1950

1951

do

do

36

157

26

T16 RS, Maine 35 115 150

T16 RS, Maine 1954 95 10 85 95

T16 R5, Maine 1952 do.- 114 70 44 114

Colton, N.Y. 1946 British 360 174 173

Columbia.

North Elba, N.Y 1947 do 525 260 262 522

' Recorded in the East prior to liberation.

' Fuparia.

o

ta

H

H

O
TABLE 3 .—^Liberations against the oriental moth (Cnidocampaflavescens (Wlkn.))

Species Origin Wliere liberated Year Where collected Number

shipped

Living

insects

released,

total

1930

1930

1930

1929

1930

1930

1930

1930

1930

1930

1930

1929

1930

1930

1930

1930

1917-20

6,395

7,129

5,680

2,221

5,848

6,011

4,947

7,076

4,983

5,865

6,854

2,326

5,146

4,457

6,171

5,063

?

6, 395

Cambridge, Mass

Chelsea, Mass

do

do

do -

do

IlllldoIIIIIIIIIII

do....

do

do_...

do.. .-

do -.

do

7,129

5,680

Dorchester, Mass

IlllldoIIIIIIIIIIIIIIIII

Everett, Mass

2,221

5,848

6,011

4, 947

Medford, Mass 7,076

4, 983

5,865

5, 854

2,326

do..

do

6,146

4,457

6, 171

5,063

( )

a

o

a

a

a

o

o

§

Known to be established.

* Small numbers.
PARASITES AND PREDATORS OF FOREST INSECTS 9

Coleophora laricella (Hbn.)

larch casebearer

Both Canada and the United States imported parasites of the larch

casebearer, Coleophora laricella (Hbn.), from Europe during the early

thirties (table 4). Two valuable parasites, Agathis pumüus (Ratz.)

and Chrysocharis laricinellae (Ratz.), have become widely established

in both countries and very high parasitization, particularly by Agathis,

has been recorded. Agathis spread rapidly in Michigan after being

recolonized there, and it was hberated in a newly discovered infestation of the casebearer in Idaho during 1960.

Dendroctonusfrontalis Ziium.

southern pine beetle

In 1892-93, A. D. Hopkins, while studying in Germany, noted that

a predaceous clerid beetle, Thanasimusformicarius (L.), was abundant

and appeared to be valuable in controUing bark beetles. He accordingly arranged for the shipment of 6,098 adult beetles and larvae to

this country. When they arrived, 2,200 were released in the forests of

West Virginia against the southern pine beetle Dendroctonusfrontalis

Zimm. There is no record that any were ever recovered. This

shipment was the first attempt to import a natural enemy of a forest

insect into this country.

Diprion frutetorum (F.)

Diprion frutetorum (F.) is a European species of sawfly that was

first discovered in this country m 1932. In 1941 it was abundant at

Southington, Conn., and at that time cocoons were heavily parasitized

by Dahlbominus fuscipennis (Zett.). This is interesting because no

direct releases of the parasites were made to control D. jruUtorurn.

The nearest release point was Orange, Conn., some 25 miles to the

southwest where this parasite was hberated m 1936 against Diprum

hercyniae (Htg.). Canada has liberated several species agamst V.

Jniietorum in southern Ontario.

Diprion hercyniae (Htg.)

European spruce sawfly

The program of parasite release against the European spruce

sawfly, Diprion hercyniae (Htg.), in Canada ranks as one oí tue

greatest efforts of its kind that has ever been made agamst a iorœt

mseot (table 5). The sawfly outbreak was termmated about 1939

by an accidentaUy mtroduced polyhedrosis vu-us disease. IJtíore

then very large numbers of parasites were released. By iar tne mosc

common of these was a smaU gregarious cocoon parasite, Dahlbommus

fuscipennis (Zett.). This species proved to be easüy propagated m
TABLE 4.—^Liberations against the larch casebearer, Coleophora larieella (Hbn.)

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

cf cf ? 9 Total

*Ágaíhis pumilus {Ratz.)_ Europe

do

Woodstock, Conn

Calder, Idaho

1934

1960

1960

1960

1960

1960

1936

1933

1934

1932

1937

1937

1950

1934

1934

1934

1953

1953

1935

1936

1933

1936

1932

1933

1935

1935

1935

1933

Europe

Rhode Island.

do

do

do

do

Europe

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

do

806

1,000

600

500

1,025

500

72

234

850

78

1,796

833

1,215

718

634

908

"3,"98Ô"

1,650

1,140

2,136

800

848

1,512

4,038

1,775

1,050

264 642 806

768

404 Rocky Point, Idaho

St. Joe, Idaho 419 Thorn Creek, Idaho.,..

do

Bremen, Maine

Sydney, Maine

do

Lynfield, Mass

Ann Arbor, Mich. (E.

S. George Preserve).

Ann Arbor, Mich.

(Mud Lake Swamp).

Elmwood, Mich..

Berlin, N.H

464

14 18 32

156

350

26

882

336

600

52

893

386

860

78

1,776

722

1 215

218

234

403

"'i,"96Ö'

885

560

1,022

300

369

168

2,001

1,100

500

300

506

"'2,"56Ö"

629

690

1,042

500

479

1,344

2,037

675

718 Saranao, N.Y

*Chrysocharis laricinellae

(Rate.).

Sharon, Vt

Rhinelander, Wis

Three Lakes, Wis

Woodstock, Conn

Bremen, Maine

Sydney, Maine

908

173

174

3,960

1,514

1,140

Lunenburg, Mass

do

Winchester, Maas

Berlin, N.H..

2, 064

800

848

1,612

Lempster, N.H

Saranao, N.Y

4,038

1,775

1.025
Dicladocerus westwoodi

Westw.

Horogenes îaricineîîae

(Strobl.).

Horogenes nana (Grav.) do.

...do.

...do.

Tupper Lake, N.Y.

Sharon, Vt

Khinelander, Wis..

Three Lakes, Wis.

Lee, Mass

Woodstock, Conn.

Melrose, Mass

Sydney, Maine

Berlin, N.H

do

Saranac, N.Y

Sharon, Vt

Lee, Mass

1936

1935

1953

1953

1936

1934

1932

1934

1933

1934

1934

1934

1936

do 2,001 973 1,003 1,976

do 4,022 2,016 2,006 4,022

do 221

do 222

do 231 158 73 231

do 580 275 305 580

do 26 26 26

do 700 200 498 698

do 100 100 100

do 700 200 500 700

do 700 200 500 700

do 545 245 300 545

do 231 158 73 231

*Known to be established.

>

ido>oo

«I §
TABLE 5.—Liberations against the European spruce sawfly, Diprion hereyniae (Htg.)

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

d'cf 9 Î Total

*Apte8Ís basizonia

OJrav.).

Europe Allagasli, Maine

Appleton, Maine

Castle Hill, Maine

Cyr, Maine

1941

1943

1940

1942

1943

1941

1942

1942

1941

1940

1940

1941

1941

1940

1942

1942

1942

1942

1942

1939

1940

1940

1935

1942

1941

1939

1939

1940

1940

1939

1938

Lab. reared

do

do

do

do

do

do

do-.

do

do

do..

do

do

do

do

do.

do

dodo

do.

do

do

Europe

Lab. reared

do

do

do

do

do

do

Europe

> 5, 630

2,646

' 1, 000

' 6, 430

50

' 5, 630

767

1,000

' 6, 630

1,200

' 1, 000

' 5, 630

' 5, 630

51

' 6, 430

' 6, 430

' 6, 430

2,646

6, 430

400

1,067

'919

3,930

' 6, 430

' 5, 630

617

238

3,000

697

522

35

5 630

2 646

Ï 000

6,430

50 Grafton, Maine

Harpswell, Maine

Harrison, Maine

Jefferson, Maine

Kibbie, Maine

Mt. Desert, Maine

" Mt." Ka'tahdiii^ "M^ne"

Parlin Pond, Maine

Tl R6, Maine

T2 R13, Maine

T2 R13, Maine

T5 R9, Maine

T6 R7, Maine

T9 R5, Maine

T9 R5, Maine

T9 R5, Maine

T14 R6, Maine..

T15 Rll, Maine

T34, Maine

5 630

767

1 000

5 630

1,200

1, 000

5, 630

5 630

51

6,430

6, 430

6,430

2 646

6,430

380

460 607 1,067

919

1,580 2,350 3,930

6, 430

5,630

617

238

Jaffrey, N.H

RensselaerviUe, N.Y...

Shandaken, N.Y

Jacksonville, Vt

Marlboro, Vt

3 000

298 299 597

522

35 Wilmington, Vt 5 30
Apíeais subguttata

(Grav.).

*DahUtominu8 fuscipetif

g nis (Zett.). â Delomerisia japónica • Cush.»

Exenierus ahruptorius

(Thnb.).

do

do

Oregon

Europe .-

Dublin, N.H

Wilnaington, Vt

Natcliaug, Conn

Tupper Lake, N.Y

Ashland, Maine

Days Grant, Maine

Friendship, Maine

"TIÏ RÍ6," M"ato¿ri"II

Square Lake, Maine

T9 R5, Maine—

T9 R5, Maine

T15 RIO, Maine

T16 RIO, Maine

Westfield, Mass

do

Dublin, N.H

iiiiidor.iiiiiiiiiiiii

Temple, N.H.

"TVp'per"Lake,"Ñ"Y."II

Lincoln, Vt

Marlboroj VÍmiIIIII

IlllldoIIIIIIIIIIIIIII

WUmingtcn, Vt

do

1938

1938 «

1936

1937

1936

1937

1937

1939

1939

1939

1939

1937

1939

1939

1937

1937

1939

1939

1938

1938

1939

1938

1938

1986

1938

1938

1938

1939

1939

1938

1939

do.

do.

m

Oregon.

do.

do.

Europe,

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

do.

220

6S

«

348

SOO

330

1,400

2,468

2,000

2,340

2,343

5,000

2,256

4,800

4,800

2,600

2,800

815

1,000

3,270

3,130

3,420

3,148

3,202

1,490

3,200

3,200

3,200

1,800

2,580

3,200

40, 800

110

19

268

245"

110

46

m

SO

500

85

See footnotes at end of table.
TABLE 5.—^Liberations against the European spruce sawfly, Diprion hercyniae (Htg.)—Continued

Species Origin Wliere liberated Year Where

collected

Number

shipped

Living insects released

d'cf Î 9 Total

Exenierus amictorius

(Panz.).

Europe

. do

do

do

dodoOregon

Europe

dodo- -

do

Ashland, Maine

T9 R5, Maine

T14 R6, Maine

Dublin, N.H

1937

1940

1935

1938

1940

1940

1939

1940

1940

1940

1940

1940

1938

1939

1939

1935

1938

1935

1935

1936

1936

1936

1938

1939

1939

1939

1938

1939

1939

1939

1939

Europe

IlllldoIIIIIII

do

do

do

dodo -

do

do

do

Nebraska

Europe:

IlllldoIIIIIII

dodo:

Oregon

do

do

do

do

Europe

IlIIIdoII-Ido

do

do

do

do

do

100

281

2,696

51

173

340

400

102

450

397

246

64

37

120

50

253

144

290

325

17

237

413

163

600

65

25

35

116

474

125

350

25

75

75

21

65

121

72

206

2,529

30

108

219

97

281

2,604

51

173

Jacksonville, Vt

Marlboro Vt

340

315

Exenierus claripennis

Thorns.

T9 R5, Maine

Shandaken, N.Y

Dover, Vt

n

200

174

82

38

91

250

223

164

26

102

450

397

Exenterus pini Gush

Exenierus tricolor RomanJacksonville, Vt

do

Dublin N H

246

64

37

Marlboro Vt 119

Exenterus sp

Ischnojoppa luiecUor

(Thnb.).

Itoplectis 4-cingulaius

ffrov.).'

Lamachua eques (Htg.)..-

Lamachus marginaius

BrÍBchke

'TTÍ R6,"Maïnê"IIIII

Wilmington, Vt- -

47

92 158 253

136

174

125

1

100

65

37

200

19

116

200

16

137

348

126

400

43

i 290

Orange, Conn—'.

do

Petersham, Mass

Tupper Lake, N.Y

Wilmington, Vt

IlllldoIIIIIIIIIIIIIII

Westfleld, Mass

Dublin, N.H

325

17

237

413

163

600

62

25

35

Marlboro Vt 115

Wilmington, Vt. 309

Lamachus sp

iydella Bp

Hi

.do - 148 205 348

tt^
Monodontomerus denHpes

(Daim.). , , Olesícampe ratzeburgi

(Tsohek).

Ihorocera (?) hamata

A. & W.

Spaihimeigenia spinigera

Towns.

Sturmia sp

do

do

Nebraska--

do

Europe

Wilmington, Vt

Marlboro, Vt--

Wilmington, Vt.

Jacksonville, Vt

Westfleld, Mass,

Jacksonville, Vt,

Marlboro, Vt-_,

Known to be established.

1 Parasitized cocoons.

' In 1935 Canada supplied 30,000 Dahlbominus fuscipennis for

release in Maine. The next year about 500,000 were made

available, and the parasite was liberated at a number of points

throughout New England and New York. Further shipments

were received in 1937. By 1938 the States of Maine, New

Hampshire, Vermont, and New York began propagating and

liberating Dahlbominus on such a large scale that it is hardly

necessary to list all liberation points. Virtually all spruce stands,

showing even light infestations, were colonized in the next few

years. Total liberations of this species may be summarized as

follows:

1939

1938

1940

1939

1940

1939

do

do—,

Nebraska-,

do--.

do—.

Europe

182

295

394

572

400

8

82

274

195

92

'Ï95'

298

197

172

257

394

39

572

392

dumber

Staie liberated

Maine 226, 173, 000

New Hampshire - 5, 410, 000

Vermont 15, 540, 000

New York.. 10,420,000

Massachusetts _- 140, 000

Connecticut 90, 000

Total.-- _ 257,773,000

' Recorded in the East prior to liberation.

>

O

d

a)

a

o

wœ

O

ta

o

wm

o
16 XJ.S. DEPT. OF AGRICULTUEB HANDBOOK NO. 226

the laboratory, and large rearing programs were conducted in both

Canada and the united States, where the sawfly also was a pest.

Well over a billion individuals were hberated.

In addition to Dahlbominus another cocoon parasite, Aptesis hasizonia (Grav.), and five larval parasites became established. The

larval parasites are Exenterus claripennis Thorns., E. amictoriiis

(Panz.), E. vellicatus Cush., E. tricolor Roman, and Drino (Prosturmia)

bohémica Mesn. The most effective of the above species have been

Dahlbominus fuscipennis and E. claripennis mider conditions of high

outbreak density, and E. vellicaius and Drino bohémica at lower

densities.

Canada sent substantial numbers of all imported parasites of the

sawfly to the United States for release in this country, and large

numbers of Dahlbominus were propagated and released by several

State organizations. This species became generally established.

Since the sawfly outbreak was terminated by disease even earher in

the United States than in Canada, no concerted effort was made to

recover most of the other species released. It is very likely, though,

that those that were established in New Brunswick are now present

in Maine.

Diprion similis (Htg.)

introduced pine sawfly

No Uberations have been made against the introduced pine sawfly,

Diprion simuis (Htg.), in the United States; but it is commonly

parasitized in this country hy Monodontomenis dentipes Dahn., one

of its European enemies. Undoubtedly the latter was introduced with

its host from Europe. M. dentipes is considered an important parasite

of Diprion polytomum (Htg.), as well as D. similis in Europe.

Galerucella xanthotnelaena (Sehr.)

elm leaf beetle

No egg parasites of the ehn leaf beetle, QaleruceUa xanthomelaena

(Sehr.), are known to be established in this country in spite of the fact

that large numbers of Tetrastichus galerucae (Ponsc.) [formerly caUed

T. xanthomelaenae (Rond.)] and Tetrastichus sp. from Europe have

been released in the East, and large numbers of Tetrastichus sp. from

Japan have been released in California (table 6). Attempts to introduce a tachinid parasite, Erynniopsis rondani Towns., in the East

were also unsuccessful, but this species took hold rapidly in California,

and in recent years several good colonies have been collected there

and released throughout the United States. No concerted efforts

have yet been made to recover Erynniopsis or TelraMiekus spp. in the

East, however, and their estâbhdiment is a possibility. A native

pupal parasite, T. brevistigma Gahan, which is common in the East,

was successfully established in California.
PARASITES AND PREDATORS OF FOREST INSECTS 17

Gossyparia spuria (Mod.)

European elm scale

Three species of parasites that attack the European elm scale,

Oossyparia spuria (Mod.), in Europe have been imported into this

country and released in California. This work, which was done by

the State of California, has been summarized by S. E. Flanders in the

Journal of Economic Entomology 45:1078-79, 1952.

Shipments of host material were sent from Germany (1908), Italy

(1939), and France (1948-53). The parasites obtained were

Coceophagus insidiator (Dalm.), THehomasthus cyanifrons (Dalm.),

and Siicroterys sp. O. insidiator (Dalm.) is the only one which was

recovered and that only at Kedlands, but parasitization at the point

of release was 85 percent 2 years after colonization. C. P. Clausen

has written ' that the scale has been adequately controlled by

Coceophagus in several localities.

Hemerocampa leucostigma (J. E. Smith)

white-marked tussock moth

One of the parasites, Apanteles solitañus (Katz.), introduced against

the satin moth, and two of the parasites, CompsUura concinnata

(Meig.) and Ooencyrtus kuwanai (How.), introduced agaiast the

gypsy moth, are easily reared on the white-marked tussock moth,

Hemerocampa leucostigma (J. E. Smith), in the laboratory. They

have, therefore, been released against this host in a few locations

(table 7). CompsUura and Apanteles are frequently found parasitizing

tussock moth larvae in the field in New England, but there is no

record of Ooencyrtus having been recovered from H. leucostigma eggs

in Washington, D.C., or Iffinois where special liberations were made

against that host.

Hemerocampa pseudotsugata McD.

Douglas-fir tussock moth

Two imported gypsy moth enemies, Galosoma sycophanta (L.)

and CompsUura concinnata (Meig.), and a polyphagous pupal parasite.

Pimpla turioneUae L., originally reared from the European pine shoot

moth, were released agamst fiie Douglas-fir tussock moth, Hemerocampa pseudotsugata McD., in the West (table 8). They have not

been recovered, but no concerted effort has been made to do so.

' Peraonal communication.
TABLE 6 —Liberations against the elm leaf beetle, GaleruceUa xanthomelaena (Sehr.)

Species Origin Where Hberated Year Where collected Number

shipped

Living

insects

released,

total

Erynniopsis rondani Europe 1957

1957

1957

1955

1956

1957

1957

1939

1935

1958

1936

1934

1955

1958

1958

1944

1955

1956

1934

1934

1934

1932

1933

1932

1932

1908

1932

1933

1925

1908

1033

California

do

do...

do

11IIIdo--II—III

638

638

638

15

60

1,600

1,650

31

371

760

55

510

150

325

150

130

500

785

40

' 25, 000

' 15, 000

3,400

3,605

3,000

1,000

7

1,200

2,000

620

eoo

4, 000

638

Towns.»

do ____

Massachusetts

638

638

15

-..do

-.-do

Castle Crag, Calif

Manteca, Calif

60

1, 600

do 1,650

31

California

Europe

221

650

Assonet, Mass 52

Woburn, Mass

CaUfornia

do -

do

IlllldoII—IIIIIII

510

College Park, Md

Reno, Nev .

115

200

do

Forty-Fort, Pa

100

4

Alexandria, Va 387

Warwick, Va . 737

40

Fresno, Calif

Massachusetts

do

Gahan Orangevale, Calif

Port Costa, Calif

Visalia, Calif.

' 25, 000

» 15, 000

Fresno, Calif 3 400

(Fonsc.) and TetrasUchus do

do—

do

do

do

do

do

do

do

3,605

galerucae). Greenwich, Conn

Washington, D.C

do

do

Blackstone, Masa

Cambridge, Mass

Kingston, Maas

1,000

?

1,200

2,000

620

600

4, 000
Tetrastichus sp. Japan.

Melrose, Mass

do

Taunton, Mass

Wakefield, Mass

Woburn, Mass

do

Newfleld, N.H

Dunellen, N.J

New Brunswick, N.J_.

Plainfield, N.J.

Rutherford, N.J

Ithaca, N.Y

Saratoga Springs, N.Y.

Cincinnati, Ohio

Dayton, Ohio

Arlington, Va

Braddock Heights, Va.

Richmond, Va

Central California

1908

1911

1933

1925

1933

1935

1932

1935

1908

1935

1932

1908

1932

1932

1933

1932

1932

1933

1934, 1936,

1938

do_

do_

do_

dodo_

do.

dodododo.

dododo.

do_

do_

do_

do.,

.do.

1,200 1,200

843 643

1, 000 1,000

55 55

2,000 2,000

825 825

600 600

1,260 1,260

7 7

224 50

1,000 1,000

7 7

1,000 1,000

3,000 3,000

2,000 2,000

760 760

750 750

2,000 2,000

4,000 14, 000

Known to be established.

' A small colony was also liberated in Virginia in 1925.

> Approximate.

TABLE 7.—^Liberations against the white-marked tussock moth, Hemerocampa leucostigma (J. E. Smith)

Species Origin Where liberated Year Where collected Number

shipped

Living

insects

released,

total

Apanteles soliiarius (Ratz.) Europe 1929

1910

1917

1921

1921

Lab. reared

New England

Lab. reared

do

do

1,200

437

5,000

12, 000

20, 000

1,200

115

6,000

12, 000

20, 000

Compsilura concinnata

(Meig.).

Ooencyrtus kuwanai (How.) .

do Washington, D.C

do

Washington, D.C
20 U-S. DEPT. OF AGEICULTÜBB HANDBOOK NO. 226

Hemileuca oUvUte CMl.

range caterpillar

Three of the imported enemies of the gypsy moth, which have become established in the East, were liberated against the range caterpillar, Hemileuca oliviae Ckll., in Arizona and New Mexico between

1913 and 1929, but none of them became established (table 9).

Heterarthrus nemoratus (Fall.) {=PhyllotOTna nenwrata Fall.)

birch leaf-mining sawfly

The birch leaf-mining sawfly, Heterarthrus nemoratus (Fall.), is a

European species which was recorded from Nova Scotia as early as

1905. In 1926 it appeared abimdantly in Maine. A nimiber of parasites were shipped to this country from central Europe between 1930

and 1934. Two of them, Ohrysocharis UmdneUae (Ratz.) and Phanomeris phyüotomxie Mues., became established (table 10). The host

has been of practically no economic importance since that time, however, and no attempt has been made to determine the importance of

the parasite in recent years.

Malacosoma disstria Hbn.

forest tent caterpillar

Two imported parasites were released against the forest tent caterpillar, Malacosoma disstria Hbn., at Cass Lake, Minn., in 1937 ; a colony

of 4,711 (1,726 d'd' and 2,985 99) Oompsilura concinnafa (Meig.)

and a colony of 2,985 Pimpla turionellae (L.). CompsUura is known

to attack M. disstria in the East. Pimpla is a polyphagous pupal

parasite that was originally reared from European pine snoot moth

pupae from Europe; but it attacks a wide variety of hosts, among them

M. disstria, under laboratory conditions. No recoveries were made

following these liberations.

Malacosomafragilis (Stretch)

great basin tent caterpillar

Oompsilura condmiata (Meig.) and Calosoma sycophanta (L.) have

been collected in the East and liberated against Malacosoma spp. in

the West. A colony of 590 Calosoma was sent to AJhambra, Calif.,

in 1918 for release against either Malacosoma fragilis (Stretch) or M.

pluviale Dyar. In 1929, 1,000 Calosoma were released on the Santa

Fe (N. Mex.) National Forest and 300 were released in Alamogordo,

N. Mex. In 1942, 1,000 Calosoma and 800 CompsUura puparia were

released on the Santa Fe National Forest. Neither species has been

recovered.
TABLE 8.—-Liberations against the Douglas-fir tussock moth, Hemerocampa pseudotsugata McD.

Species Origin Where liberated Year Where collected Number

shipped

Living

insects

released,

total

Calosoma sycophania (L.) Kurope Northport, Wash, 1930

1937

1937

New England

do

Lab. reared

1,200

1,000

2,000

1, 002

Compsilura concinnata do

do

985

(Meig.).

Pimpla turionellae (L.) do 1,968

TABLE 9.—Liberations against the range caterpillar, Hemileuca oliviae Ckll.

Species Origin Where liberated Year Where collected Number

shipped

Living

insects

released,

total

Calosoma sycophanta (L.) Europe Cimarrón, N. Mex*

Koehler, N. Mex

Koehler and Maxwell,

N. Mex.

Raton, N. Mex - _

1928

1913

1914

1929

1915

1929

1914

1915

1914

New England

do --

do -

IlllldoIIIIIIIIIII

Lab. reared

100

30

1,400

2,000

2, 000

• 2, 200

1 3, 150

' 2, 005

?

100

do

30

1,400

1 199

Compsilura concinnata Tempe, Ariz 1

CMeig.). Raton, N. Mex 700+

400+

?

Roawell, N. Mex

do

Maxwell, N. Mex

' Puparia.
to

ts3

TABLE 10 —Liberations against tlie birch leaf-mining sa wñy, Heterarthrus nemoratus (Fall.)

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

rf'cf ? ? Total

, Bar Harbor, Maine 1933

1933

1935

1931

1935

1935

1934

1933

1935

1933

1933

1935

1931

1932

1934

1935

1931

Europe

do

-.-.do—

....do

.—do

—-do

-—do—

—.do—

-.-.do

--do -

—.dodo

do

...-do

—do

-...do -

do

1,345

1,350

1,940

63

567

1,700

450

404

125

275

200

1,116

88

28

210

50

12

603

600

500

15

130

500

150

79

25

742

750

1,440

48

290

1,200

300

325

100

1, 345

(Ratz.).

....do... .

1, 350

Eustis, Maine . 1, 940

Strong, Maine

North Andover, Mass..

Stark, N.H.

63

420

1 700

Bolton, Vt , 450

Bar Harbor, Maine.....

Eustis, Maine

404

....do........

..-.doi.;.':!.^.

125

Bar Harbor, Maine

Bethel, Maine

273

200

488

60

ÜÖ"

628

28

28

100

50

12

1, 116

Scambus foliae (Cush.)...

North Conway, N.H...

Keene, N.Y

Eustis, Maine . ._

88

28

210

50

Tramsema pedellu -

(Hlmgr.).

North Conway, N.H... 12

!»

O

H

O

g

H

O

O

M

§

• Known to be established.
PAKASITBS AND PREDATORS OF FOREST INSECTS 23

Miscellaneous releases of Calosoma

The large carabid, Calosoma sycophanta (L.), which was successfully

introduced into New England from Europe against the gypsy moth, is

known as a rather omnivorous feeder, although it prefers gypsy moth

larvae and pupae. Because of this, small colonies of Calosoma beetles

were collected in New England and colonized against a variety of hosts

widely distributed throughout the United States. Calosoma has never

been recovered in any of these release areas, but a list of host species

is presented here because of possible interest (table 11).

Miscellaneous releases of Compsilura

The tachinid ñj, Compsilura concinnata (Meig.), which was successfully established m this country as a parasite of the gypsy and browntail moths, also attacks the imported satin moth and more than

200 native hosts. Because of its exceptionally polyphagous habits, it

was collected in New England and released against a few crop and

fruit insects in some rather widely separated areas. It has not been

recovered from these hosts, but the liberations are listed in table 12.

They may prove of interest in their distribution even if Compsilura

fails to attack the host against which the release was made.

Neodiprion abietis (Harr.)

balsam fir sawfly

In 1943 the imported European spruce sawfly cocoon parasite,

DaUhomimis juseipennis (Zett.), was Uberated against the balsam fir

sawfly, Neodiprion abietis (Harr.), in the New Harbor, Maine, area by

entomologists of the State of Maine. A note in the Biennial Report

of the Forest Commissioner for the State of Maine for 1943-44, page

37, states that cocoons collected at the liberation point were heaviy

parasitized.

Neodiprion fulviceps (Cress.)

In 1941 a shipment of 200,000 DaUhominus fuscipennis (Zett.)

was sent to Deadwood, S. Dak., for release in the Harney National

Forest against Neodiprion fulviceps (Cress.). The parasite has not

been recovered, but no special collections have been made for that

piupose.

Neodiprion lecontei (Fitch)

red-headed pine sawfly

When large numbers of the parasite DaUhominus fuscipennis

(Zett.) were being reared for liberation against the European spruce

sawfly from 1939 to 1941, a considerable number were also liberated

against the red-headed pine sawfly, Neodiprim lecontei (titch).

About 580,000 were released on pine plantations of the Tennessee

Valley Authority. Three hundred thousand of these were hberated

in northwestern Alabama near WUson Dam in 1940. The remainder

were put out in the same general area in western Tennessee m 1941.
M

!*>.

TABLE 11 .—Miscellaneous releases of Calosoma sycophanta (L.)

te

Host species Locality Year

Calosoma beetles

Shipped Released

AUophila pometaria (Harr.).

ArUicarsia sp

Carpocapaa pomonella (L.). -

Datana integerrima G. & R._.

Epilachnn varivestris Mills. _-

Hyphanlria cunea (Dru.)

Phryganidia califomica Pack.

Rhopobata naevana (Hbn.). _

Blowing Rock, N.C

Linville, N.O -

Gainesville, Fla

Bentonville, Ark

Grand Junction, Colo

Greenwood and Winchester, Va

Farmington, Del

Birnaingham, Ala

Sacramento, Calif

Alhambra, Calif

Seaview, Wash

1921

1921

1915

1915

1915

1915

1917

1921

1913

1918

1919

?

166

400

12

158

16

457

963

600

35

50

30

590

45
TABLE 12.—Miscellaneous releases of Compsilura coneinnata (Meig.)

I

t1

a

s

I

01

§

Host species Locality Year

Compsilura

Puparia

shipped

Adults

released

CoUas phiîodice eurytheme Bdv

Harrisina briUians B. and McD

Hyphantria cunea (Dru.)

Laphygma frugiperda (J. E. Smith)

Tempe, Ariz

Sacramento, Calif

Riverside, Calif

San Dieeo County, Calif

Portland, Oreg

Gainesville, Fla

1918

1923

1924

1952

1933

1915

1916

10, 635

1,000

1,000

9,000

896

2,125

7

?

Î

4,790

74

?

?

to
26 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

That same year 200,000 were released on the Maniatee National

Forest in Michigan. In New York 530,000 were released in 20

towns in and around the Adirondack region. In 1946 a shipment

from Canada was released near Cadillac, Mich. Few attempts were

made to recover the parasite at any of these liberation points, and

the only place where the parasite is known to have been established

is near Cadillac, Mich.

In 1932 a colony of 120 Sturmia inconspicua (Meig.) (=Drin,o

bohémica Mesn.) was liberated at Mont Alto, Pa. It was not known

whether this European species would attack Neodiprion lecontei

or not, but it seemed a possibility. No recoveries were made.

Neodiprion pratti pratti (Dyar)

During the summer of 1959 the Division of Forestry of Virginia

recovered Dahlbominus fuseipennis (Zett.) from Neodiprion pratti

pratti (Dyar) cocoons collected in several counties in the central

part of the State. More than 23,000 Dahlbominus were reared in

the laboratory and colonized at a number of points that season, and

more than a million were reared and released in 1960. No releases

of this parasite were made previously in Virginia, so it must have

spread naturally from New Jersey or Marjdand.

Neodiprion pratti partidoxicus Ross

A colony of 6,000 parasites, Exentems airuptorivs (Thnb.), supplied by Canada from European collections, was liberated against

the sawfly Neodiprion pratti paradomcus Ross at Ashby, Mass., in

1940. The parasite has not been recovered.

Three smaU colonies of Aptesis basizonia (Grav.) reared from

European stock at New Haven, Conn., were released against a

Neodiprion sp. in 1940. It is believed that this was the same species

later described by Koss (Forest Sei. 1: 196-209, 1955) as N. pratti

paradoxicus. Liberations were as follows: Groton, Mass., 200;

Greenwood, N.Y., 103; Pittsford, Vt., 100.

E. E. McCoy of the New Jersey State Department of Agriculture

has reported that Aptesis basizonia (Grav.) has been recovered from

"paradoxicus" in New Jersey.'

The State of New Jersey has also reared DaMbominus iuscipennis

(Zett.) on a large scale for release against this pest. Since 1957,

1,863,000 have been colonized in Neodiprion pratti paradoxicus

infestations and according to Mr. McCoy a number of good field

recoveries have been made.'

Neodiprion sertifer (Geoff.)

European pine sawfly

Canada has conducted an intensive program of parasite introduction against the European pine sawfly, Neodiprion serfvfer (Geoff.),

and large colonies of several species of parasites have been released

in that country. Shipments of these species have also been sent to

' Personal communication.
PARASITES AND PREDATORS OP POKEST INSECTS 27

the United States for release (table 13). In addition, the State of

New Jersey has propagated large numbers of Dahlbomimtsfuscipennis

(Zett.) for release in N. serii/er infestations. Three species are known

to be established. Of these, Daklbominus is generally distributed

throughout the_ State, and Aptesis basizonia (Grav.) is also well

established. Limited recoveries of the larval parasite, Exenterus

abruptoñus (Thnb.), have also been made by the Northeastern Forest

Insect Laboratory. (It is worth noting in this connection that

Canada also imported a polyhedrosis virus disease of N. sertifer

from Europe and that it has been used effectively against the larvae

in both Canada and the United States.)

Neodiprion swainei Midd.

Swaine's jack-pine sawfly

Professor R. D. Shenefelt released about 100,000 DaUbominus

fuscipennis (Zett.) against Swaine'sjack-pine sawfly, Neodiprion swainei

Midd., in Iowa County, Wis., in 1953.

Nygmia phaeorrhoea (Donov.)

brown-tail moth

Large scale importations of parasites and predators of the browntaU moth, Nygmia phaeorrhoea (Donov.), were conducted concurrently

with those of the gypsy moth from 1905-11. The work has been

described in detau in U.S. Department of AgricultiKe Technical

Bulletin 86 (1929), and the summary of liberations made through 1927

(table 14) is taken from table 34 in that pubUcation. In order to

expedite dispersion into the Maritime Provinces, the Canadian

entomologists collected Calosoma sycophanta (L.) and Meteorus versicolor (Wesm.) adults and OompsUura concinnata (Meig.) puparia in

New England for reoolonization during 1912-15. Releases of two

additional species in the United States are recorded in table 15.

Nine beneficial species became established in this country. Two of

them, Oompsilura concinnata (Meig.) and Exorista larvarum (L.),

are perhaps more properly Hsted, however, as gypsy moth parasites

inasmuch as they are even more important enemies of that pest. The

most effective of the brown-tail moth parasites are Tovmsendiellomyia

nidicola (Tns.) [now called Alsomyia nidicola Tns.], Apanteles lacteicolor Vier., and Meteorus versicolor (Wesm.).

Porthetria dispar (L.)

gypsy moth

The work done to estabMsh predators and parasites of the gypsy

moth, PoHhetria dispar (L.), in the United States is so weU known

that it need not be described m detaU here. Large scale importations

from Europe were made from 1905 to 1914 and from 1922 to 19ád.

Several species were also obtamed from Japan during the early penod.

Table 16 summarizing this work through 1927 is taken from U.b.U.A.
TABLE 13.—Liberations against the European pine sawflj, Neodiprion sertifîr (Geoff.)

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

d'd' ? 9 Total

*Aptesia basiionia

(ferav.).

Europe

do

Oregon

Europe

do

do

do

East Orange, N.J.

Millburn, N.J.. _-

1940

1940

1940

1940

1940

1941

1941

1941

1941

1941

1941

1941

1941

1941

1941

Lab. reared...

do

do

do..

do

1,000

317

952

1,000

1,000

400

4,800

2,654

3,190

6,400

6,400

75

112

83

94

13

1,000

317

do

Lamington, N.J

Feapack, N.J

952

1 000

1 000

*DMbominus fusciven-

» nis (Zett.).

Delomerista japónica

Gush.«

Exenten« abrupioriiu

(Thnb.).

New Jersey m

Ohio

Far Hills, N.J Oregon %no

Europe

IlllldoIIIIIII

do

do

do

do

do.

do

do..

Gladstone, N.J

Harding, N.J

612

1,148

253

3,088

1,954

57

64

13

24

3

4,188

1,506

2,937

3,312

4,446

18

48

70

70

10

4,800

2 654

Lamin^on, N.J

do

Millburn, N.J

3,190

6,400

6 400

75

|*(Panz.).

Iiehnojoppa luteaior

(Thnb.).

Lamachua eques (Htg.)...

do

do.

Gladstone, N.J

Harding, N.J

112

83

94

13

to

00

*Known to be established.

' In 1939 the U.S. Department of Agriculture liberated 480,000,

and from 1939-45 the State of New Jersey liberated 12,644,000

Daklbaminus in iV". aertifer infestations scattered throughout the

State.

' In 1939 the U.S. Department of Agriculture supplied the

Ohio Agricultural Experiment Station with 390,000 Dahlbominua

which were liberated m N, sertifer infestations in Brainbridge and

Clyde, Ohio.

• Recorded in the East prior to liberation.

O

o
TABLE 14.—Foreign enemies of Nygmia phaeorrhoea (Donov.) liberated in North America through 1927 ^

Species

Individuals

of foreign

stock liberated

By reproduction from

foreign stock

Subsequently liberated—

From New

England field

collections

Total

enemies

liberated >SO>

e

o

ai

o

*Xponíe¡es lacUicolor Vier

*Carahus auratus L

*Carcelia laxifrans Vill

*Compsilura coneinnata (Meig.)

*Eupteromaluí nidulaní (Thoms.).

*Ekorista larvarum (L.)

*MeteoTua verticolor (Wesm.)

*Monodontomenis aereus Wlkr

*Tovm8endiellomyia nidicola (Tns.)

Number

55, 000

478

9,742

25, 134

7

42, 152

3,113

15, 541

3,500

Number

255, 245

530, 000

122, 625

7,887

Number

310, 245

478

9,742

147, 759

530, 000

42, 152

11, 000

15, 541

3,500

*Known to be established.

1 From U.S.D.A. Tech. Bui. 86.

TABLE 15.—^Liberations against the brown-tail moth in the United States, 1928-59

Species Origin Where liberated Year Where collected Number

sliipped

Living

insecte

released,

total

Blondelia piniariae (Htg.).-- 1929

1929

1929

12, 169

9,335

120

12, 169

9, 335

-do ---

Nashua, N.H do

Carcelia rutila Meig. York, Maine do 120

CO
TABLE 16.—Foreign enemies of Porthetria dispar (L.) liberated in Nortli America through 1927 ^

Species

*Anastatus disparis Ruschka

AparUeles Uparidis (Bouché)

*Apanteles melanoacelus (Ratz.)

Äpanteles porthetriae Mues.

AparUeles solitaTius (Ratz.) '

Blepharipoda scuteüata (R.~D.)

Bl&ndelia nigripes (Fall.)

Brachymeria intermedia (Nees)

Brhchyfneria obecurata (Walk.)

Caloaoma chinense Klrby

CaloSoma inquisitor (L.)

Caloaoma teticulatum (Fab.)

*Caloaoma aycophanta (L.)

Carabua arvenis Fab. (.=arcensia Herbst.)

*Carabus auratus L.ä*

Carabus glabratus Payk

Carahus violaceus L

*Carabus nemoralia Müll.

Careelia laxifrons Vill

Carcelia separata Rond

*Compsilura concinnata (Melg.) *

CrossocosTnia ñavoscutellata Schiner

Crossocosmia sericariae (Rond.)

*Exorista larvarum (L.)

Bxorista segregata (Rond.)

Masicera eyloatica (Fall.)

Meteorus japonicue Ashm.

Individuals

of foreign

stock

liberated

Number

138, 680

76, 702

23, 476

12, 065

22, 546

11,097

10, 692

20, 798

394

140

259

83

2,711

108

478

63

136

9,742

17, 061

25, 134

700

42, 152

9, 323

23

S

Subsequently liberated—

By reproduction from

foreign

stock

37, 370

"22,"S22'

128

27

27

By reproduction from

established

stock

132, 177

From New

England

field

collections

Number

65, 505, 513

73, 546

63, 870

122, 625

Total

enemies

liberated

Number

65, 644, 193

114,072

155, 653

34, 587

22, 546

84, 643

10, 692

20, 798

394

268

286

110

66, 581

108

478

63

136

9,742

17, 061

147, 759

700

42, 152

9,323

23

400

03

O
Meíeorus pulchrícornis (Wesra.) - . 4

1,703

582

18, 445

12, 543

} 402

75

13, 364

471

4,568

9,420

10, 499

118 122

*Ooeneyrtzts kuwanai (How.) ... 25, 675, 884 25, 677, 687

-rafes pavida Meig : . . 582

*Parasetigena süaestria (R.-D.) . 2,278 20, 723

*PAo6o(;ompe díspom (Vier.) .... 12,543

Pimfja emminator (Y&h.) . .

75

Sturmia inconspicua Melg. {—Drino bohémica Mesn.) . 13,364

471

Tachina magnicomis (Zett.) __._.._.. 4,568

Taehinids unclassified • ... 9,420

Tachinids unclassified ^ ...... 10^499

4,650

76, 000

15

4,650

76, 000

Xylodrepd Quadripitnctattí Sehr. 100

504

lis

504

Known to be established.

I From U.S.D.A. Tech. Bui. 86, p. 142.

ä Some doubt as to this species.

* Also liberated against Nygmia phaeorrhoea (Donov.).

* Includes some multibrooded species liberated in 1906-07.

' Mostly Exorista larvarum in 1926.

* Number of foreign stock received unknown.

2

>

09

M

o

>
32 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

Technical Bulletin 86. Table 17 summarizes work done since 1927.

Nine parasites and two predators were established in this country.

Two of the parasites are egg parasites. Both are well distributed and

often abundant throughout the infested areas. One of these, Ooencyrtus kwwanai (How.), is from Japan and is the only parasite established

from that country. The other, Anagtatus disparis Kusohka, is from

central Europe.

Four taohmid larval parasites are also well distributed and abundant. They are Oompsihira concinnata (Meig.), Exorista larvarum

(L.), Parasetigena sUvestris (R.-D.), and Blepharípoda scuteUata

(R.-D.). One hymenopterous larval parasite, Apanteles meUmoscdus

(Ratz.), can also be placed in this category. Two other larval parasites, Phobocampe disparis (Vier.) and Monodontomerus aereus Wlkr.,

are rather scarce and unimportant. One predator, a large carabid,

Calosoma sycophanta (L.), is very abimdant and destructive. The

other, Carabus auratus L., is of no practical importance.

Generally speaking there is an excellent sequence of insect enemies

operating against the gypsy moth in this country, but the almost

universal use of insecticides when defoUation occiffs compUcates an

evaluation of the long-term results of this work.

Pristiphora erichsonii (Htg.)

larch sawfly

An ichneumonid parasite, Mesoleius tenthredinis Morley, was imported from England and successfully colonized against the larch

sawfly, Pristiphora erichsonii (Htg.), in Canada in 1910-11. For

many years it was an extremely effective parasite, and it spread naturally throughout the Lake States west to Montana and east as far as

Pennsylvania. A few small colonies were also released in the United

States, but it is doubtful if they affected spread of the parasite.

According to information supplied by Prof. S. A. Graham, in 1911

a small colony of Mesoleius was released on Grand Island near Munising, Mich. Dr. H. J. MacAloney of the U.S. Forest Service reports

that eight colonies were liberated in 1953 in two general areas in

northern Minnesota. In 1929, 14 specimens were released in Rindge,

N.H., and in 1931 two small colonies of 42 and 35 specimens, respectively, were released at Lunenburg, Mass.

In 1953 Prof. R. D. Shenefelt released about 100,000 DaMbominus

fuscipennis against the larch sawfly in Douglas County, Wis., and

about the same number in Burnett County, Wis. Two colonies were

also released in New York State, one at Kent in 1940, and one at

Sanford in 1939. Each of these contained 10,000 parasites.

Rhyacionia buoliana (Schiff.)

European pine shoot moth

Both the United States and Canada have undertaken extensive

programs of parasite introduction against the European pine shoot

moth, Rhyacionia buoliana (Schiff.), and large numbers of parasites

from England and the Continent have been released in this country
TABLE 17.—Liberations against the gypsy moth, Porthetria dispo r (L.), 1928-59

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

cf cf 9 9 Total

*Anastatus disparts

Ruschka

Europe

do

do

do,.

do

do

do-_

do

New England ^

Duxbury, Mass

Ashburnham, Mass

w

Thompson, Conn

Middleborough, Mass..

Bow, N.H _

(.)

1932

1929

1928

1932

1931

1930

1928

1929

1928

1929

1930

1931

1930

1930

1934

1934

1930

1933

1930

1928

1930

1930

1934

1934

(')

New England.

do

do

2,880

352

3,336

2,880

(Bouché).

*Apantelea melanoscelus 275

1,522

77

1,814

352

3,336

w

(£-D.) .

Europe

IlllldoIIIIIII

do

do

dodo

do

do -

New Englanddo

do

do

do

do

do

do

do

do

do

do

163

360

7

53

12, 177

1,454

10, 981

895

304

800

800

168

193

916

142

750

"716

1,400

'35

100

100

163

360

7

Nashua, N.H 53

Blondelia piniariae Htg. Amherst, N.H

Rye, N.H---

12, 177

1,454

Cranston, R.I _ 10, 981

Brachymeria intermedia Dighton, Mass

Plymouth, Mass

Barkhamstead, Conn.--

Wethersfield, Conn

Woodstock, Conn

Limerick, Maine

Agawam, Mass---

Belchertown, Mass

Bernardston, Mass

Carlisle, Mass

895

(Nees). 304

*Calosoma sycophanta 800

(L.) 800

168

193

916

142

750

716

Deerfield, Mass

Dighton, Mass

Erving, Mass

1, 400

35

100

Gill, Mass 100

See footnotes at end of table.

CO
TABLE 17,-—Liberations igainst the gypsy moth, Porthetria dispar (L.) 1928-59—Continued

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

cfcf Î 9 Total

*Calosoma »ycophanta

(L.)—Continued.

Europe

do

do

Greenfield, Mass

Hatfleld, Mass

Holden, Mass

1934

1930

1928

1929

1929

1934

1929

1929

1933

1928

1928

1928

1928

1930

1930

1930

1930

1929

1931

1932

1930

1928

1929

1928

1929

1931

1928

1930

1929

1928

1928

New England.

do

do.

do

do

do

do

do

do

do

do

do

do

do.

do

do

do

Europe

IlllldoIIIIIII

do.

do

do

-.—do

do

do

do

do

do

do

New England.

100

950

276

5,000

66

90

5,000

6,000

127

150

557

506

364

700

1,000

1,423

1,000

3,870

1,669

6

2,034

1,052

30

484

120

250

2,300

3,978

66

16

' 3, 184

100

950

149 126 275

Mendon, Mass

Middleton, Mass

Montague, Mass

Northampton, Mass

Princeton, Mass

Ware, Mass

5,000

66

90

5,000

5, 000

127

Westford, Mass

Windham, N.H

East Greenwich, R.I...

Wakefield, R.I

Brattleboro, Vt

Granby, Vt...

75

345

238

198

75

212

268

166

150

567

506

364

700

1 000 Westfleld, Vt 1 423

Carcélia separata Rond.-- Westminster, Vt

Chelmsford, Mass

Middleborough, Mass..

Saugus, Mass

1, 000

3,870

1, 659

6

Bow, N.H 2 034 Nashua, N.H 1 062

*Compñlura eoneinnata

do....

Boxford, Mass

Chelmsford, Mass

Middleborough, Mass..

Rowley, Mass

30

484

120

250

Bow, N.H 3,978

66

16

3,184

Nashua, N.H.

Rye, N.H

Westerly, R.I
Emeatia 'fip

*Bxorista larvarum (L.)-.

*Ooencyrtus kuwanai

(How.).

*Para3etigena aihiestris

(E.-D.).

do..

do..

Japan...

Europe.

*Phobocampe disvaris

(Vier.).

Phobocampe sp-.

do-

.do_

Bow, N.H

Canton, Conn

Thompson, Conn..

Chelmsford, Mass.

Saugus, Mass

Bow, N.H

Manchester, N.H_.

Nashua, N.H

do

Easton, Mass

Foxboro, Mass

Mansfield, Mass..-

Norton, Mass

Thompson, Conn..

Union, Conn

Acton, Mass

do

Boxford, Mass

Chelmsford, Mass.

Duxbury, Mass..

Holden, Mass

Kehoboth, Mass.

do

Sherborn, Mass..

Sudbury, Mass.-,

Biddeford, Maine..

Sebago, Maine

Hillsdaie, N.Y

Attleboro, Mass

Baxnstable, Mass..

Dighton, Mass

.doStow, Mass..

1930

1941

1932

1929

1932

1930

1930

1928

1929

1928

1928

1928

1928

1933

1941

1932

1933

1928

1929

1933

1930

1931

1932

1930

1931

1930

1933

1942

1930

1929

1931

1932

1930

Europe

do

do

do

do

do

do

do

do

Lab. reared.,

do

do ,

do

Europe

do

do

do

do

do

do

do

do

do

do

do

do

dodo

do

do

do

do

do

5

65

1,112

1,530

21

23, 221

3,323

2,370

149

224, 000

144, 000

56, 000

88, 000

2,880

199

16, 782

2,880

9,622

»6,538

11, 429

» 10, 318

200

129

11, 875

22, 017

1,197

1,830

« 3, 650

» 10, 946

2,576

112

1,862

500

194

58

73

5

23 42 65

912

1,530

21

23, 221

3,323

2, 370

149

224, 000

144, 000

56, 000

88, 000

1,200

100

5,759

1,200

4,137

1,680

99

11, 023

1,680

5,485

2,880

199

16, 782

2,880

9,622

6,322 6,107 11,429

6,224

""i,"3Ô5"

200

129

7,075

15, 793

1,197

530

200

129

11, 875

22, 017

1,197

1,830

300

45

1,120

2,276

67

742

2,576

112

1,862

500

126

23

23

68

35

50

194

58

73

See footnotes at end of table.

CO
TABLE 17.—Liberations against the gypsy moth, Porthetria dispar (L.) 1928-5f —Continued

Origin Where liberated Year Where

collected

Number

shipped

Living insects released

cfcf ? Î Total

Sturmia inconspicua

Meig. {=Drino bohémica Mesn.).

Europe

do

Beverly, Mass

Lunenburg, Mass

do

Middleborough, Mass..

1928

1928

1931

1931

1929

1930

1930

1928

1928

1929

1933

1929

1932

1930

1928

Europe

do

do

do

do

do

do

do

dodo_

do

do -

do

do

do

13, 314

11,598

980

1,495

245

624

492

116

2,963

24

16

7

3

21

1,004

13, 314

11 598

980

1,495

245

do

Bow, N.H

624

492

Nashua, N.H 116

Middleton, R.I_

Chelmsford, Mass

Melrose, Mass

2, 963

ZenilUa libalrix (Panz.)-- 24

16

7

"BOW, N.HI"IIIIIIIII

3

21

Nashua, N.H 1,004

05

Known to be established.

Ï More than 65 million Anastatus disparis were colonized

throughout New England from 1908 through 1927. About 8^

million were released from 1928 through 1932. It is unnecessary

to list individual releases for such a widespread program, but

numbers by States from 1928 to 1932 are given in order to complete

the early record published in bulletin 86.

State

Number

liberated

2 From 1928 through 1933 large numbers of Blepharipoda scutellata puparia obtained from gypsy moth pupal collections were

recolonized throughout the infested area. Since the species is

widely dispersed, individual liberation points have not been given.

A summary by States follows:

Liberation Puparia

State points liberated

Connecticut 9 20,847

Massachusetts 53 118,030

Vermont 13 32,957

Connecticut 600, 000

Maine 2, 818,000

Massachusetts — - 4, 431,000

New Hampshire __ 620, 000

Rhode Island 50, 000

Total 8, 419, 000

Total _ - 171,834

In addition 300 adults were liberated at Hillsdale, N.Y., in 1942,

and 5,594 adults were liberated at Westerly, R.I., in 1928.

3 Larvae.

* Puparia.

5 Parasitized host larvae.

O

>

»IM

a

d

!?

O

a

o

o

M
PARASITES AND PHBDATOES OF FOREST INSECTS 37

smd Canada (table 18). At least four species have become estabUshed

although one of them, the larval parasite Orgilus obscurator (Nees)'

was undoubtedly present in the United States prior to Hberation'

The other, Temducha interruptor Grav., a larval parasite, and two

pupal parasites, Tetrastichus turionum (Htg.) and Pimpla turionellae

(L.), have never been recovered in substantial numbers, although

both are important parasites at times in Europe. P. turionellae has

been recovered in Canada but not in the United States. The shoot

moth is especially difficult to control with insecticides, and it is

unfortunate that its parasitic enemies have been of such little consequence in this country.

Rhyacionia frustraría bushnelli (Busck)

pine tip moth

Tip moths are very serious pests of pine on the extensive man-made

pine forests of the Nebraska National Forest. In 1925 pine tips

infested with Rhyacioniajrustrana (Comst.) were collected in Virginia

and taken to Nebraska in order to obtain parasites for liberation

against B. Jrustrana bushnelli (Busck). Several species were liberated

that year, and work was supplemented by some further collections

from Massachusetts in 1926 (table 19). Gampoplex frustranae Cush.

took hold immediately and within 5 years appeared to be bringing the

pest under control. Unfortunately about this time another variety of

tip moth, R. frustrana neomexicana Dyar, increased very rapidly.

Campoplex fnistranae does not parasitize neomexieana, so conditions

were not substantially improved.

Setora (Miresa) nitens Wlk.

nettle grub

In 1932, at the request of the Commonwealth Institute of Biological

Control, 5,000 oriental moth cocoons, some of which were parasitized

by Chaetexorista javana B. & B., were sent to Ceylon from Melrose

Highlands, Mass. It was hoped that this parasite could be recovered

from these cocoons for release against the nettle grub, Setora {Miresa)

nitens Wlk. About 500 Chaetexorista adults were obtained.

Stilpnotia salicis (L.)

satin moth

The satin moth, Stilpnotia salicis (L.), was first discovered in this

country at Medford, Mass., in 1920. Thatsame year it was also found

at Westminster in British Columbia, and 2 years later at BeUingham.

Wash.

Several species of parasites that attack the gypsy and brown-tail

moths also attack the satin moth, so it is not surprising that some of

them were found attacking Stilpnotia salicis when it was discovered

in New England. Between 1927 and 1934 several more species were
TABLE 18.-—Liberations against the European pine shoot moth, Uhyacionia huoliana (Schiff.)

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

<?<? 9 9 Total

Actia nudihasis Stein Europe

do

do

do

do

do

Branford. Conn

Easton, Conn."

1935

1934

1932

1937

1937

1937

1937

1935

1932

1936

1933

1937

1932

1933

1934

1937

1932

1933

1937

1933

1933

1932

1933

1933

1937

1937

1937

Europe

iiiiidoiiiiri:

do

do

do

do

do

do

do

do

do

do

do

do

do

do..

do

do

do

do

do

do

do

do

294

157

49

96

60

24

101

18

39

65

43

504

1,666

16, 000

7,000

10, 000

1,603

16, 000

12, 380

16, 000

7,500

1,162

15, 000

14, 000

10. onn

285

Maiden, Mass 18

41

29

12

45

7

7

21

13

168

633

5,000

33

55

31

12

56

10

32

34

30

334

1,033

10, 000

49

96

60

24

101

17

Campoplex borealis

(Zett.).

Campoplex muUvyinctus

Gray.

Campoplex mutabilis

(Holmg.).

Meriden, Conn

do __

do. ..

Chester, N.J...

CarrmopUx ruHfemm

Crtioms.).

Campoplex sp

Buffalo, N.Y

Branford, Conn

Fairfield, Conn

Brookline, Mass

Chester, N.J

do -.

39

66

43

602

1,666

16, 000

6,878

9 674

Copidosoma genimlatum

(Dalm.).

Branford, Conn

Easton, Conn

Fairfield, Conn

Hamden, Conn

do..

Meriden, Conn

North Greenwich,

Conn.

Brookline, Mass

Hingham, Mass

Wakefield, Mass..

Winchester, Mass

Boonton, N.J

352 1,151 1,503

16, 000

12 380

16, 000

7, 500

214 948 1,162

16 000

14' 000

9,000

18,500

9.000

Chester, N.J do 19,500 1 New Vernon, N.J do 10,000

CO

00
Ephiiütes ruücoUie

(Grav.).

Lypha dubia (Fall.)-

Macrocenlrus sp.

Orgilus obicurator (Nees).

...do.

Carmel, N.Y

do

North Castle, N.Y-__.

do

Oyster Bay, N.Y

do ----

Plainview, N.Y

Rockland Lake, N.Y_.

Southeast, N.Y

Syosaet, N.Y

Washington, N.Y

Bristol, Pa

Fairfleld, Conn_

Buffalo, N.Y

Lewiston, N.Y

Niagara Falls, N.Y...

Branford, Conn

Easton, Conn

Hamden, Conn

Brookline, Mass

Jericho, N.Y

Meriden, Conn

Chester, N.J. _-.

Branford, Conn

Easton, Conn

Fairfleld, Conn

Meriden, Conn

Brookline, Mass

Wakefield, Mass

Winchester, Mass

Ooeana County, Mich

Boonton, N.J

New Vernon, N.J

Buffalo, N.Y

Jericho, N.Y

Lewiston, N.Y

Niagara Falls, N.Y._.

Southeast, N.Y

Syosset, N.Y _..

1932

1933

1934

1937

1932

1933

1934

1933

1932

1937

1933

1933

1936

1935

1935

1936

1935

1934

1938

1933

1934

1937

1937

1932

1934

1935

1936

1931

1933

1933

1959

1937

1936

1935

1932

1935

1935

1932

1936

_do_-_

.do-_-

.do.--

_do.--

_do.--

_do.--

_do.--

_do„-

_do.-_

_do_-

_do_-_

.do_—

.do.-,

-do...

-do.--

-do...

-do...

-do.--

.do...

.do...

.do-.-

-do--

-do...

.do.-,

.do...

.do.-,

.do.-

-do-..

-do...

-do.--

-do...

.do.-,

-do--

-do--

-do--

-do-..

.do.--

-do--

-do.--

1,100 400 700

15 000

3' 000

10, 000

1,100 400 700

15, 000

10, 000

15, 000

1,100 400 700

10, 000

15, 000

15, 000

554 243 311

38 11 24

948 75

304 67 203

74

200

41

2 39

113 45 68

500

252 139 113

150 50 100

100 100

92 34 55

150 46 104

278

72 32 40

10 10

110 110

100 33 67

170 86 65

372 185 184

28 14 11

50 38

799 166 577

799 166 577

100 100

320 152 167
TABLE 18.—Liberations against the European pine shoot moth, Rhyadonia buoliana (Schiff.)—Contiaued

Pimpla lurioneUae (L.)

Origin

Europe-.

PrülomeTut vulnerator..

(Panz.).

*Temelucha interruptor

Grav.

.-.do.

....do.

Wtiere liberated

Branford, Conn.

do

Fairfield, Conn..

do

do

Meriden, Conn.,

.do.

Newtown, Conn

Boonton, N.J

Hackettstown, N.J.

New Vemon, N.J...

Oakland, N.J

Buffalo, N.Y

Lewiston, N.Y

Niagara Falls, N.Y.

North Caatle, N.Y..

.do.

Roslyn, N.Y

Syosset, N.Y....

do... _

Meriden, Conn..

Branford, Conn.

do -.

do.

Easton, Conn...

do L-...

Fairfield, Conn.,

do

Hamden, Conn.

Meriden, Conn.,

Year

1935

1936

1935

1936

1937

1936

1937

1936

1937

1938

1935

1938

1935

193S

1935

1936

1938

193S

1936

1938

1937

1932

1936

1936

1934

1936

1935

1936

1936

1936

Where

collected

Lab. reared.

do

do

do

do

do

do

do

.....do

do

do

do

Europe

do

do

Lab. reared..

do

do

do ,

do

Europe

do

do

do

do

do

do

do

do

do

Number

shipped

242

664

524

378

1,000

500

1,000

58

500

400

65

400

10

35

60

432

1,000

1,000

150

1,000

64

200

450

454

171

490

143

850

450

600

Living insects released

cfc?

100

317

324

126

250

1

IS

20

181

34

SO

100

67

89

58

97

60

100

9 9

142

324

200

250

1,000

260

1,000

68

500

400

65

400

8

19

26

248

1,000

1,000

150

1,000

30

200

400

354

101

400

85

747

400

600
* Tetrastiches turionum

(Htg.).

Trichogramma evavf

escens Westw.

do

-do.

Brookline, Mass 1931

Hingham, Mass 1931

Wakefleld, Mass 1933

Winchester, Mass 1933

Boonton, N.J 1937

New Vernon, N.J 1936

Buffalo, N.Y 1935

Jericho, N.Y 1932

Lewiston, N.Y 1935

North Castle, N.Y 1936

Eoslyn, N.Y 1936

Southeast, N.Y 1932

Syosset, N.Y 1936

Branford. Conn

Ëaston, Conn

1933

1934

Fairfield, Conn 1935

North Greenwich, Conn_ 1936

Brookline, Mass 1933

Newton, Mass 1933

Chester, N.J 1937

New Vemon, N.J 1935

Bedford, N.Y 1933

Matineoock, N.Y. 1933

North Castle, N.Y 1933

Plainview, N.Y 1934

Fairfield, Conn 1937

Meriden, Conn 1936

do 1937

Chester, N.J 1937

do

do

do

do

do

do

do

do

do

do

do

do

do

.-.do

do

do

do

do

.—do

Lab. reared.

Europe......

IllldoIIIIi:

do

do

Lab. reared..

do

do

do

390

275

10

140

80

985

272

218

7,172

986

1,000

225

1,000

20, 000

2,000

10, 675

42, 000

20, 000

20, 000

5,000

10, 000

20, 000

20, 000

22, 000

5,600

35, 000

4,500

40, 000

35, 000

155

76

27"

485

120

"i,"43r

474

495

"""söö"

235

200

10

140

48

495

132

218

5,144

496

500

225

500

6,250 13, 750

4,000 6,000

6,150

9,450

13, 300

12, 550

390

275

10

140

75

980

252

218

6,575

970

995

225

1, 000'

20, 000

2,000

10, 475

42, 000

20, 000

20, 000

5,000

10, 000

19, 950

19, 450

22, 000

5,600

35, 000

4,500

40, 000

35, 000

"Known to be established.

13O

>i m
0^

60

TABLE 19.—Liberations against the pine tip moth, Bhyaeionia frustrana bushndli (Busck)

Species Origin Where Uberated Year Where collected Number

shipped

Living

insects

released,

total

*Campoplexfrustranae Cush.. Virginia Halsey, Nebr 1925

1925

1926

1926

1925

1925

1925

1925

1925

1925

1925

369

1,437

?

Î

13

254

?

Î

Î

7

15

369

Eurytoma pini Bugbee

Bxeristes comstockii (Cress.)-

Olyptavaripes Cress

Habrocytus thyridoptengis

Massachusetts

do

Virginia

IlllldoIIIIIIIIIIIIIIIII

IlllldoIIIIIIIIIIIIIIIII

Massachusetts

do

Virginia

1,437

13 How.

HaÜichelía rhyacioniae Gahan.

Hyssopus rhyacioniae Gahan.

Hyssopus ihymua Gir

lAxophaga mediocrii Aid

lAxophaqa plúmbea Aid

Temelucha epagogea (Cush.).

do

do

do...

do

do..

do

IlllldoIIIIIIIIIIIIIIIII

IlllldoIIIIIIIIIIIIIIIII

IlllldoIIIIIIIIIIIIIIIII

do

do

do

do

do..

do...

254

15

Known to be established.

' "A few."

O

a

o

%

3

g

H

O
PARASITES AND PREDATORS OP FOREST INSECTS 43

imported from central Europe, and all the promising species were also

colomzed m the Pacific Northwest (table 20). Those known to be

established in the Northeast are Apanteks solitarius (Ratz.)

Eupteromalm nidulans Thorns., CompsUura concinnata (Meig.), ana

Exaiista larmrum (L.). The first three have become established in the

West and, in addition, A/eieoros versicolor (Wesm.) has also become

established there. This is rather surprising because it has never

become established in the East.

In both areas parasites are generally credited with holding the pest

at very low levels. Aparddes solitarms seems to be the most important species in both areas. Meteoros is very important in the West;

CompsUura and Eupteromalus in the East.
TABLE 20.—Liberations against the satin moth, Stilpnotia solids (L.)

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

cfcf V 9 Total

*Apanteles solitariua

(Rat«.).

Europe.

do

do

do

Chatham, Mass..

Danvers, Mass

do

Ipswich, Mass

Natick, Mass

1932

1927

1927

1929

1928

1929

1927

1928

1932

1930

1930

1929

1930

1933

1930

1927

1939

1930

1928

1930

1932

1933

1928

1928

1930

1930

1929

1929

1935

1931

1931

Europe

Lab. reared-..

do..

do

do

do

do

do..

New England.

do

do

do

do

Europe

IlllldoIIIIIII

do..

do

do

do

do

do

New England.

do

}....do-

}....do

do

do

do

472

443

' 21, 250

113, 659

1,389

200

205

1,150

335

100

100

200

30

420

5,000

860

3,188

1,813

1,115

2,758

9

300

415

13

2 2, 350

• 1, 850

»253

230

' 1, 000

472

266 177 443

853 536 1,389

do

Yarmouth, Mass

Nashua, N.H

200

76

525

13

129

625

234

205

1,150

Kent, Wash 247

Orting, Wash 100

Puyafiup, Wash.

Seattle, Wash ...

100

198

Sumner, Wash

New Haven, Conn

Augusta, Maine

Danvers, Mass

30

Careelia gnava Meig 351

5,000

860

3,188

Yarmouth, Mass

Nashua, N.H...

1,813

1, 115

Newmarket, N.H_

Kent, Wash

2,768

' 3

do

Nashua, N.H

92

*Compmlura concinnata 405

(Meig.). Newmarket, N.H

Bellevue, Wash

Kent, Wash

13

Í 1,057

Puyallup, Wash

Tacoma, Wash

Puyallup, Wash

Sumner, Wash

do

I 300

143

59

?

:^
f Eupteromalus nidulans a Thorns.

Exorisia lanarum (L.).

*Meteoru> veràeolor

(Wesm.).

do-

.do.

...do

do

do

do

Silvana, Wash

Sumner, Wash

Tenino, Wash

Anacostes, Wash

Centralia, Wash

Everett, Wash

Kent, Wash

Tacoma, Wash

Olympia, Wash

Topsham, Maine

Natick, Mass

Nashua, N.H

Newmarket, N.H

Sumner, Wash

New Haven, Conn

Portland, Maine

Skowhegan, Maine

South Hai'pswell,

Maine.

Topsham, Maine

Chatham, Mass

Harwiehport, Mass

Ipswich, Mass

Natick, Mass

South Yarmouth, Mass.

Taunton, Mass

Concord, N.H...

Conway, N.H

East lungston, N.H...

Newmarket, N.H.

Bellingham, Wash

Centralia, Wash

Kent, Wash

do

Stanwood, Wash

Sumner, Wash

1932

1933

1933

1933

1934

1932

1933

1934

1932

1931

1932

1933

1933

1929

1928

1930

1933

1933

1933

1933

1934

1933

1932

1932

1929

1929

1933

1932

1933

1932

1931

1930

1934

1934

1932

1933

1934

1933

.do

.do

.do

do-,

do_.

do_.

do..

do..

do-.

do-.

doEurope.

do..

do..

do..

do..

do..

do..

do..

do..

do..

do..

do..

...do..

...do..

do..

do..

...do-,

...do..

...do..

...do..

do..

do..

dodo..

dodo996

340

100

045

443

885

650

500

656

000

755

180

309

247

120

000

500

500

500

000

500

600

900

531

636

500

874

400

085

000

840

000

000

200

250

000

250

1,258 1,242

181 819

220

400

273

380

500

1,258

192 682

267

391

818

1,609

504 696

See footnotes at end of table.
TABLE 20.—Liberations against the satin moth, StUpnotia salicis (L.)—Continued

Species Origin Where liberated Year Where

collected

Number

shipped

Living insects released

cfcf 9 9 Total

Pales -pavida Meig Europe

do

Washington

Europe

Naticls, Mass 1929

1932

1928

1932

1934

1933

1934

1933

1933

1932

1933

1934

1934

1934

1934

1934

193S

1929

1932

Europe

IlllldoIIIIIII

do

do

do

do

do

do.

do

do

do

do

do.......

do

do

Washington

Europe

61

33

757

56

300

500

500

500

500

200

150

500

500

500

432

500

104

75

75

61

Lexington, Mass

Nashua, N.H

33

757

Kent, Wash 46

Rogos unicolor (Wesm.) _ . New Haven, Conn

Portland, Conn

South Harpswell,

Maine.

Topaham, Maine

South Yarmouth, MassTaunton, Mass

Concord, N.H

293

500

500

500

500

200

150

Woonsoolcet, R.I

BelUngham, Wash

Kent, Wash. . . ...

500

497 ' 492

Olympia, Wash..

Stanwood, Wash

Waterbury, Conn

Natiok, Mass .

430

457

TacMnomyia similis

.(WittO^ • ... ,

ZmiUia Kbatrix (Panz.)--

104

75

Kent, Wash 2

Known to be established. ' Parasitized host larvae. • Fuparia.
INDIVIDUAL SPECIES SUCCESSFULLY

ESTABLISHED IN THE

UNITED STATES

Agathis pumilus (Ratz.) (Braconidae)

Agathis pumuus (Ratz.) was introduced into Canada and the United

States from Europe in the late twenties and early thirties along with

several other larch casebearer parasites. It was rare in England but

fairly important in southern France and Austria, where it parasitized

12 to 20 percent of the host popidation. It became established readily

andjs now common throughout the range of the casebearer in North

America except in a newly discovered infestation near St. Maries,

Idaho. It was colonized there in 1960.

Collections hj A. R. Graham in southern Ontario in 1957 showed

average parasitism by Agathis of 41 to 67 percent. In 1960 limited

collections in Connecticut, Massachusetts, and Pennsylvania showed

equally high parasitism in some isolated stands of larch. Parasitism

in generaUy infested natural stands in Nova Scotia ran about 25

percent.

Agathis has shown remarkable power of dispersion. For example,

in 1937 it was liberated by Prof. S. A. Graham near Ann Arbor, Mich.,

and by 1951 had spread north of the Straits of Mackinac. In 1950

it was released near Iron River, Mich., by F. E. Webb, and in 1953

it was recovered 50 mUes to the east.

Agathis has one generation a year. The winter is spent as a first

instar in overwintering casebearer larvae. The parasite does not develop beyond the first instar until the host larva ties its case down

preparatory to pupation. The parasitized larva never pupates. The

parasite completes development and pupates within the host; adults

emerge during late June or early July.

Anastatus disparis Ruschlia CEupelnúdae)

.Anastatus disparis Ruschka, an important e^ parasite of the gjTsy

moth, occurs in both Europe and Japan. It was first introduced into

this country in 1906 and first recovered in 1910. It can be readüy

propagated and easUy collected. More than 65 milhon parasitized

eggs have been distributed throughout the infested region, üetaoled

records of parasitism were kept for many years at regular coUecfaon

points, most of which were somewhat north of Boston, Mass. ine

highest average percentage parasitism was 28 percent m 19¿l-¿¿.

This declined to 6 percent 5 years later, but undoubtedly these hgi^es

are closely associated with Host population fluctuations and otner

"inlate years Anastatus has been scarce in southern Massachusetts

and Connecticut, and another egg parasite, OoencyHus kuwanai, has
48 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

been abundant in that area. It is, therefore, generally believed that

Anastatus is more important in the northern range, and Ooenq/rtus

more important in the southern range of the gypsy moth in this

covmtry. This needs confirmation. AnasiattK is often important in

southern climates, such as that in Spain, so it can hardly be considered

"a more northern species."

Observations by D. E. Parker indicate that there is little conflict

between the two parasites. Nevertheless, both species are rarely, if

ever, abundant in one location.

Anastatus has one generation a year. The winter is spent as a fullgrown larva in the nost's egg. Adults emerge in June and July.

Females oviposit in newly laid egg clusters, and often Aiiastatus can

be seen ovipositing in fresh eggs, as they are being laid by the female

gypsy moth. Larval development requires about 2 weeks, and the

full-grown larva remains inactive until the following spring.

Apúnteles lacteicolor Vier. (Braconidae)

Apartides ladeicolor Vier, became established in this country in 1908.

It is now generally distributed throughout the area infested by the

brown-tail moth. In 1925, parasitism of overwintering larvae in 28

towns averaged almost 7 percent. In some places it was as high as

25 percent in single webs.

Apanteles lacteicolor overwinters in brown-taü moth larvae. Adults

emerge in the spring, and two summer generations may develop on

other hosts, including the gypsy moth. Females oviposit within the

body cavity of the host. The mature larva emerges to spin its silken

cocoon in the brown-tail web or on tree trunks or leaves. Hibernating

brown-tail moth larvae are attacked in August.

Apúnteles melunoscelus (Ratz.) (Braconidae)

Apanteles melanoscelus (Katz.), a European parasite of the gypsy

moth, is at times very important locally, particularly in the southern

range of the host in Europe. Large numbers were sent to this coimtry

in 1911 and 1912. The parasite became established almost immediately and was recolonized widely. Average parasitism is rather

low, but almost every year it is abimdant locaUy m widely separated

areas. This parasite is very heavily attacked by hyperparasites,

which must cut down its effectiveness tremendously.

Apanteles melanoscelus is primarily a solitary parasite of the gypsy

moth, although it occasionaUy attadks other hosts. It completes two

generations annually upon the gypsy moth. Winter is spent in the

cocoon stage. Adults appear soon after gypsy moth larvae hatch in

May, and eggs are laid internally in first mstar host larvae. Parasite

larvae complete development, and emerge from second and third

instar larvae to form cocoons. Adults emerge 7 or 8 days later, and

third-stage host larvae are attacked by second generation females of

the parasite. The second generation larvae form cocoons which

overwinter, and since they are usually in exposed places they are

subject to attack by hyperparasites over a very long period.
PARASITES AND PREDATORS OP FOREST INSECTS 49

Apúnteles solitarius (Ratz.) (Braconidae)

Avanteles sólüañus (Ratz.) was sucessfully introduced into NewEngland from Europe against the satin moth in 1927. It became

established immediately, and dispersed rapidly throughout the

infested area. In 1932 it was colonized in the State of Washington

where it took hold equally well. A. solitarius has been an exceptionally effective parasite in both areas. For example, parasitism of

hibernating larvae at a number of collection points in New England

in 1933 averaged between 15 and 67 percent, and in Washington in

1935 from 0 to 55 percent. Host populations were very much reduced

in New England in 1935, and parasitism averaged 6 to 11 percent.

These figures represent a minimum, however, for they are based upon

hibernating host larvae, and A. solitarius also kills some larvae before

they enter hibernation and others after they emerge in the spring.

The species is a solitaiy internal parasite, which has been reared

from several hosts in the laboratory but only from the satin moth in

the field.

The seasonal history of Apanteles solitarius is somewhat complex

because it may overwinter as a first instar larva within the host larva

or within a cocoon. From both places adults emerge and attack small

satin moth larvae in the spring. Cocoons formed at this time may

overwinter, or they may produce adults that attack the small hibernating host larvae in late summer.

Aphidecta obliterata (L.) (Coccinellidae)

Aphidecta obliterata (L.) is widely distributed in northern and central

Europe where it feeds on a number of Chermids. Canadian entomologists have imported it since 1951 for release against the balsam

woolly aphid, Chermes piceae, and colonies have been sent to the

United States for release in the Pacific Northwest and North Carolina.

Although it had completed a summer generation the year of release

in New Brunswick, it apparently cannot survive Canadian winters.

It is included here because it has recently completed a summer generation in North Carolina and may have a better chance of survival there

than in New Brunswick.

Little is known regarding the effectiveness of Aphidecta, but it is

reported as feeding commonly on Chermes on twigs in Europe, and a

predator with this feeding habit might be of considerable value here.

Aphidecta has one generation a year. The winter is spent as an

adult in the ground. Eggs are deposited on the needles or trunks of

infested trees. Both adults and larvae attack all stages of the balsam

woolly aphid except the first instar larvae.

Aphidoletes thompsoni Mohn (Cecidomyiidae)

Aphidoletes thompsoni Mohn is a Cecidomyiid predator of Chermes

piceae in Europe. Lar^e numbers have been collected, particularly in

Czechoslovakia, and shipped to Canada. Canadian entomologists in

turn have sent excellent colonies to the United States for release in

New England, the Pacific Northwest, and North Carolina. So far,

it has not been recovered in Canada or New England, but it has become
50 U.S. DEPT. OP AGEICULTUEE HANDBOOK NO. 228

established at several places in the Pacific Northwest. It is too early

to determine whether it has taken hold in North Carolina. Judging

from the large numbers collected in Europe, it may be of some importance as a control factor there. Since it is very small and requires Uttle

food to complete development, it might be effective at low host population levels.

In Europe there are three to five generations annually. Adults

apparently do not feed. The females lay eggs in wax masses of the

Chermes and on the bark surface. The larvae feed on Chermes eggs

and larvae. Pupation occurs in the sou within a silken cocoon.

Winter is spent as an immature larva in the soil.

Aptesis basizonia (Crav.) (Ichneumonidae)

Aptesis basisonia (Grav.) is a European parasite of sawfly cocoons,

and at times it parasitizes Neodiprion seriijer heavily. It was sent

originally to Canada for release against the European spruce sawfly,

and several colonies were made available for liberation in the United

States. These were augmented with stock reared by the U.S. Department of Agriculture and the State of Maine. Releases in this country,

as in Canada, were made against N. sertijer and Diprion hercyniae.

Aptesis has not been recovered as a parasite of the European spruce

sawfly in the united States, but small numbers were recovered from

that host in New Brunswick. It was recovered from N. sertifer at

several points in New Jersey in 1948. E. E. McCoy, New Jersey

Department of Agriculture, has also reared it from the native N.

pratti paradoxicus in New Jersey.

Aptesis completes two to four generations a year in Europe. Winter

is spent as a prepupa within the host's cocoon. Adults emerge quite

early and oviposit in overwüitering sawfly cocoons. Larvae feed

externally and one parasite develops per host. In the field, the host

larva or prepupa is paralyzed within its cocoon, but in the laboratory

this ability seems to weaken successively so that the host remains

active and is able to crush the parasite's eggs.

Blepharipoda scutellata (R.-D.) (Tachinidae)

Blepharipoda scviellata (R.-D.) is one of the most common tachinid

parasites of the gypsy moth in central Europe. It was liberated in

this country as early as 1908 and known to be established by 1911.

It was recolonized extensively, but this was probably unnecessary for

it dispersed very widely by itself. In Europe it has been recorded

from several hosts, but apparently it is restricted to the gJTisy moth

in this country. B. scutellata is one of the most abimdant gypsy moth

parasites here. In 1923, when it seemed to reach its maximum, 49

percent of the female pupae were parasitized in collections from 25

towns. Generally speaking, parasitism runs high in most areas, with

average parasitism fluctuating from 5 to 25 percent.

Blepharipoda scviellata is a large tachinid fly, which lays a microtype egg on the foliage. The ^g hatches after it is ingested, and the

parasite remains in the first instar until the host has pupated. It

then develops rapidly. The full-grown maggot emerges and drops
PARASITES AND PBEDATOES OF FOREST INSECTS 51

to the ground, where it forms its puparium and overwinters. Parasitism of female gypsy moths is considerably higher than that of males

because female caterpillars pass through an additional instar and

consequently eat considerably more foliage that may bear Blepharvpoda eggs.

Calosoma sycophanta (L.) (Carabidae)

The large, brilliant carabid beetle, Calosoma sycophanta (L.), was

iatroduced from Europe and released in this country against the

gypsy moth in 1906. The beetle became established right away, and

with further importations and recolonization soon spread throughout

most of the infested area and beyond. It feeds freely on a wide

variety of insects, especially lepidopterous larvae and pupae, but the

gypsy moth is especially suited to its taste and habits. A number of

colonies were also released throughout the South and West against a

variety of pests, but O. sycophanta has not been recovered in those

areas.

Calosoma is a conspicuous predator on gypsy moth larvae and

pupae, and it undoubtedly destroys enormous numbers annually; but,

as is true of other predators, it is difficult to estimate the actual number

of hosts killed. Collins and Holbrook found that the average number

of both larvae and pupae destroyed over a 6-year period in several

selected plots was about 25 percent. Calosoma populations buUd up

rapidly in heavy infestations but fall off rapidly when the gypsy moth

becomes scarcer. In a medium infestation in Freetown, Mass., in

1927, for instance, about 2,000 beetles were trapped per acre. The

following year when the infestation was very light, and no defoliation occurred, the number dropped to 224. Populations as high as

4,000 per acre have been recorded.

Adult Calosoma beetles live from 2 to 4 years. Winter is spent as

an adult beetle in the ground, and emergence takes place about June 1.

Eggs are laid in the ground, and the young beetle larvae soon climb

trees to feed upon caterpillars and pupae on the trunk and larger

branches. When they are fuU fed, they construct a pupal cell in Qie

soU. The adult beetle remains in the pupal cell during the winter.

Campoplex frustranae Cush. (Ichneumonidae)

Campoplex Jrustranae Cush. is a common parasite of the Nantucket

pine tip moth, Rhyacionia Jrustrana (Comst.), in the East. In 1925

it was liberated in the Nebraska National Forest against B. jrustrana

busknelli (Busck), which was severely damaging extensive young pine

plantations, particularly ponderosa and yellow pine. The parasite

became established immediately and increased very rapidly. In fact,

by 1930 parasitism had reached 82 percent at the original liberation

point, and hopes ran high that the parasite would hold the pest under

effective control. Unfortunately, another tip moth, B. Jrustrana

neomexicana Dyar, in which C. frustranae cannot develop, increased

very rapidly about this time and conditions were not substantially

improved. Parasitism of bushneUi dropped as neomexicana populations increased. It has been suggested that the parasite may have
52 U.S. DBPT. OF AGRICULTURE HANDBOOK NO. 226

become less effective on biishnelli because it deposited many of its

eggs in neomexicana.

The life history of Campoplex Jrustranae is well sjmchronized with

that of Bhyacionia frustrana bushneUi in Nebraska, where both have

two complete broods each year. The first parasites emerge after the

peak of moth emergence is passed. Young host larvae are available

when the parasites appear. Parasitized tip moth larvae complete

their development and pupate. The parasite spins its white cocoon

within the shattered pupal skin of the host. In the overwintering

brood the cocoon is not spun until spring. Since the second-brood

host larvae spin their cocoons in the sand or litter, the spring adults

emerge from host pupae formed on the ground. This is in contrast

to the situation that prevails when B. Jrustrana is parasitized in the

East. Here C. Jrustranae overwinters as a pupa within a pine bud.

Carabus spp. (Carabidae)

Carabus auratus L. and C. nemoralis MüU. were imported from

Europe in the early years to attack the gypsy and brown-taü moths.

Records regarding them are incomplete, and they have not been

recorded as feeding on these pests in this coimtry, but both species

are now widely distributed in New England. G. nemoralis was

actually recorded here prior to the gypsy moth importation program,

and as Dr. M. E. Smith notes (Ent. Soc. Wash. Proc. 61(1): 7-10.

19S9) both species may have arrived through accidental introductions.

Prof. C. O. Dirks, University of Maine, notes that both species are

present in Orono, Maine. Both species reportedly feed upon earthworms.

Carcelia laxifrons Vill. (Tachinidae)

The brown-tail moth parasite, Carcelia laxifr<ms Vill., was easily

established in New England about 1908. It has spread throughout

the infested area. Occasionally an individual collection of larvae

will be quite heavily parasitized, but the average percentage parasitism

is very low.

The only known host of Carcelia laxifrons in this country is the

brown-taU moth on which one generation is completed annually.

The winter is passed as a puparium in the soil. Adult flies emerge in

the spring and oviposit on small brown-tail moth larvae soon after

they emerge from hibernation. The maggot feeds internally completing development in the full-grown larva.

Chaetexorista javana B. & B. (Tachinidae)

Chaetexorista javana B. & B. is a large tachinid parasite of the

oriental moth. It was imported into the United States from Japan in

1929 and 1930, and several large colonies were released in the area

infested by the moth in the environs of Boston, Mass. The parasite

took hold immediately, and parasitization reached about 60 percent

in 1933 and again in 1940. Heavy mortality of parasite larvae occurs

during subzero temperatures, but the parasite increases rapidly during

milder winters.
PARASITES AND PREDATORS OP FOREST INSECTS 53

Chaetexorista javana completes only one generation annually.

Winter is spent as a second instar larva within the hibernating host

prepupa. A puparium is formed within the host cocoon. The adult

fly emerges by pushing off the cap at the end of the cocoon, which

normally serves as an exit for the adult moth. The female fly deposits

a large egg on the integument of the host larva, to which it is attached

by means of an integumental funnel. Only one parasite per host

larva develops to maturity.

Chrysocharis laricinelleie (Ratz.) (Eiilophidae)

Chrysocharis laricinellae (Ratz.) was introduced into Canada and

the United States from Europe in the late twenties and early thirties

to combat the larch casebearer, Gokophora lariceUa. The parasite

had not been reared from the casebearer in North America previously,

but it may have been present. About this time it was recovered

from the elm leaf miner, Fenusa ulmi Sund., many miles from the

closest liberation point. Studies by A. R. Graham in Canada (Ent.

Soc. Ont., 79th Rpt., 1948) showed that Chrysocharis acts as a secondary parasite of Ooleophora at times, attacking Agathis in the early

summer, and that it is not an effective parasite of the casebearer.

Chrysocharis laricinellae is also a parasite of the birch leaf-mining

sawfly, Heterarthrus nemoratus (Pall.). It was also established against

this host in New England in the early thirties. It is the most effective

parasite of Hete/rarthrus in Austria, but this host has been relatively

unimportant in New England for many years, so effectiveness of this

parasite has not been checked.

Chrysocharis laricinellae may have three generations annually in

the larch casebearer. The parasite hibernates as a full-grown larva

in the host larval case. Pupation takes place in early May, and the

emerging adult attacks the overwintermg casebearer larva. The

adults producing the second generation emerge from the middle of

June to the middle of July, and the early ones parasitize larvae already

parasitized by Agathis. Late émergents parasitize very small needle

mining larvae. Adults producing the third generation emerge from

mid-September to early October. In the birch leaf-mining sawfly,

there is one and a partial second generation annually. The winter is

spent as a full-grown larva in the mine of the host. The adult

females feed heavily on the body fluids of the sawfly larvae, and kUl

large numbers of the host.

Coccophagus insidmtor (Dalm.) (Eulophidae)

Coccophagus insidiator (Dalm.) is a Evffopean parasite successfully

introduced into California, where it has adequately controlled the

European elm scale in several localities. The same species was

apparently accidentally introduced in the East, for it was found

parasitizing the European elm scale quite heavily at Ithaca, N.Y.,

in 1924.

Coccophagus insidiator is a solitary parasite. The female is an

endoparasite of the elm scale, the male an ectoparasite of the larval

stage of the female parasite. Females oviposit in the body cavity of

the ehn scale, and development is completed in about 3 weeks. There

are three and possibly five generations annually.
54 U.S. DEPT. OF AGRICULTUBB HANDBOOK NO. 228

Compsilura concinnata (Meig.) (Tachinidae)

Compsüura concinnata (Meig.) is undoubtedly one of the best

known larval parasites of the gypsy moth. It is exceptionally

polypbagous, and attacks at least 200 lepidopterous hosts, including

the brown-tail and satin moths. Oddly enough it never appeared to

be of exceptional importance in rearing work conducted in central

Europe in the twenties and thirties; but it has been consistently

valuable in much of the area infested by the gypsy moth in New

England, achieving parasitism of 10 to as high as 50 percent. Even

higher parasitism is common in brown-tail and stain moth infestations.

It was successfully established in the Pacific Northwest against the

latter and recovered there in 1932.

Compsilura is multibrooded, completing three or four generations

each year. The winter is passed as an unmature larva within the

hibernating host larva or cocoon. Development is completed in the

spring, and the full-grown maggot emerges to form its puparium in

the duff or litter. Adult females pierce the skin of the host larva with

a sharp ovipositor and insert a first instar maggot that becomes

full grown in about 2 weeks. From one to foin: maggots may develop

in a single caterpillar.

Copidosoma geniculatum Dalm. (Encyrtidae)

Large numbers of Copidosoma geniculatum Dalm. collected in

Europe from 1932 to 1937 were released against the Em-opean pine

shoot moth in this country. It has never been recovered in the

United States, but J. L. Martin recovered it from Exotdeia dodecella L.

in southern Ontario in 1956 and 1957 (Ganad. Ent. v. 91, May 1959),

and it is believed it reached Canada on nursery stock infested by that

host. Ehyacionia buoliana and E. dodecella frequently infest the

same trees, and Arthur and Juillet * have noted that Copidosoma has

been abundant in European shipments containing both Ä. buoliana

and E. dodecella, but absent from English shipments where E. dodecella

was not present. It therefore seems likely that Copidosoma was

actually reared from Exoteleia and liberated against the wrong host.

Copidosoma geniculatum, is a tiny polyembryonic Chalcidoid which

overwinters in an immature stage with the host larva. Development

is completed about the time host larvae complete development in

early summer, and an average of about 16 parasites pupate within

each larval skin. There is one generation annually.

Cremifania nigrocellulata Cz. (Chamaemyiidae)

Cremifania nigroceUvlata Cz. is a European chamaemyiid fly

predaceous on the balsam woolly aphid throughout central Europe.

It has been released by Canadian entomologists in New Brunswick

each year since 1952, and has become established at all release points.

The predator has also been released in Oregon and recovered the

' Arthur, A. P. and Juillet, J. A. The introduced parasites of the European

pine shoot moth, RhyaeUmia bwoliana (Schiff.), with a critical evaluation of their

usefulness as control agents. Canad. Ent. 93 (4): 297-312. 1961.
PARASITES AND PREDATORS OF FOREST INSECTS 55

same season (1959). Fairly large numbers have developed on some

stem infestations in New Brunswick, but spread has been very slow.

Oremijania has up to three generations a year. The first preys upon

the spring generation of Ghermes piceae and the second and thircl on

the summer and fall generations. It overwinters as a puparium,

mostly in the ground but also on the bark. Approximately 95 percent of the first generation puparia remain in diapause during the

summer, and one-half of these continue in diapause through the

winter.

Dahlbominusfuscipennis (Zett.) (Eulophidae)

Bahlhominus iuscipennis (Zett.) is a European parasite of several

sawflies. Nearly 8 million were collected in cocoons of Neodiprion

sertifer Geofif. in Europe in 1934 and sent to Canada for release against

the European spruce sawfly. Owing to the ease with which the

species can be propagated, very large numbers have been reared and

released by Canada, the U.S. Department of Agriculture, and several

States.

Summarizing very briefly: About 226 million were released against

the European spruce sawfly in Maine, and another 30 million were

released against this same pest in New York, New Hampshire, and

Vermont. About 13 mUlion were released against Neodiprion sertifer

in New Jersey. About 1 million were released against the red-headed

pine sawfly in several States. In addition, smaller colonies have

been released against several sawflies in diJBferent States, and the

State of Virginia is currently rearing Dahlbomimis fuscipennis for

release against A^. pratti pratH.

In Europe Dahlbominusfuscipennis has been found parasitizing up

to 78 percent of the Neodiprion sertifer cocoons on the Adriatic Coast,

where soil temperatures are high and cocoons are exposed. In other

locations, such as Austria, where soil temperatures are cool and the

cocoons are usually formed in the duff, parasitism usually runs much

lower often being nil or axoimd 2 to 10 percent.

Dahlbominus fuscipennis became established quickly in Canada

and the United States, and it has been recovered at many points

throughout the colonized area. In New Brunswick it reached its

highest level in 1939, when 11 percent of the European spruce sawfly

cocoons in the southern part of the Province and 2 percent in the

northern part were parasitized.

In the United States it has been recovered in at least small numbers

from Diprion hercyniae (Htg.) in New England, Neodiprion sertifer

(Geofi.) in New Jersey, N. lecontei (Fitch) in Michigan, N. pratti

pratti Dyar in Virginia, N. pratti paradoxicus Koss and N. pini-rigidae

(Norton) in New Jersey, N. namvlus Schedl. in Wisconsin, A'', abietis

(Harr.) in Maine and D. frutetorum (P.) in Connecticut. The latter

is of interest because the nearest release point was some 25 miles

distant (Orange, Conn.), where the parasite was released against

D. hercyniae.

Percentage parasitism by DaMbominus fuscipennis has run rather

low in this coimtry, except locally. In New Jersey about 47 percent

of the Neodiprion sertifer cocoons in the top of the litter were parasitized in a representative pine plantation in 1943, and parasitization
56 U-S. DEPT. OF AGBICXILTUBE HANDBOOK NO. 226

of Diprion frutetorum averaged about 25 percent for 3 years in a small

plantation in Southington, Conn.

Dahlbominus Juscipennis is an external parasite of sawfly larvae

and pupae within their cocoons. Several eggs are laid with one

insertion of the ovipositor, and an average of 34 parasites develop in

one cocoon. Two and a partial third generation are completed

annually in southern New Brunswick. The winter is spent as a fullgrown larva or pupa within the host cocoon.

Drino (Prosturmia) bohémica Mesn. (Tachinidae)

Drino bohémica Mesn. is a European parasite of sawfly larvae,

which was successfully established in Canada against the European

spruce sawfly. It has maintained itself better than any of the other

European spruce sawfly parasites at low host population levels, and

fairly high percentages of parasitism occur when the sawfly is scarce.

It has also been reared from Pikonema olaskeTisis (Roh.) and P.

dimmockii (Cress.), and its ability to attack other hosts successfully

when Diprion hercyniae is not available is imdoubtedly important to

its survival.

Drino bohémica is probably the same species that was introduced

into the United States from central Europe as Siunnia inconspieua

(Meig.). At least a good part of that material originally came from

sawfly hosts. It was released in this coimtry against the gypsy and

brown-taü moths and also against Neodiprion lecontei. D. bohémica

persisted at one of the gypsy moth colonization sites for several years,

but apparently it failed to become permanently established in this

country at that time. It undoubtedly occurs now in northern Maine,

for it is abundant just across the border in New Brunswick and

Quebec.

Drino bohémica completes one and a partial second generation

annually in central New Brunswick. The majority overwinter as

puparia in the forest floor, but some overwinter as larvae within the

host. Most puparia are formed outside the host's cocoon. Adults

emerge the latter part of June. Females oviposit on the integument

of fifth instar larvae. The parasite maggot feeds internally usually

completing development after the host cocoon is spun.

Erynniopsis rondani Towns. (Tachinidae)

Erynniopsis rondani Towns, is a European tachinid fly parasitic on

the elm leaf beetle, particularly in southern France. Several small

colonies were imported and liberated on the East coast, but it was

never recovered. In 1939 the University of California miported 55

puparia and succeeded in liberating 31 mated females at three colonization sites. These releases resulted in establishment, and in 1940-41

parasitization reached 20 percent at one collecting point. From 1955

to 1958 collections were made in California for rdease in Arkansas,

Idaho, Nevada, Washington, D.C., and Virginia. It is not yet known

whether any of this work was successful.

Erynniopsis rondani overwinters as an immature maggot in the

hibernating ehn leaf beetle adult. The pupariimi is formed within

the dead beetle. The adult flies that emerge in the spring oviposit on
PARASITES AND PREDATORS OP FOREST INSECirS 57

beetle larvae. The parasite maggot develops internally, and there are

two summer generations. The second generation maggot remains in

the host, however, when it transforms into an adult.

Eupteromalus nidulans (Thomson) (Pteromalidae)

Very large numbers of Eupteromalus nidulans (Thomson) were

obtained from the overwintering webs of the brown-tail moth collected

in Europe from 1905-08, and the species soon became distributed

throughout the infested area in this country. It has never been

reared in substantial numbers from the brown-tail moth in the

United States, but it has proved to be a common hyperparasite of

Apúnteles and other Braconids. Fortunately, it also attacks the

overwintering larvae of StUpnotia solids. Often 20 percent of the

overwintering larvae are parasitized hj Eupteromalus, and probably

many more are kiUed for the adult parasite has the habit of puncturing

and feeding on hibernating larvae. The species was also established

in the Pacific Northwest against the satin moth.

Eupteromalus has severd generations a year. The winter is spent

as a full-grown larva in the web or hibernaculum of its host. Pupation

takes place in the spring, and the adults emerge in May and June.

The ovipositing female paralyzes the host larva and deposits her egg

on the body. The parasite larva feeds externally. The female often

feeds upon the body fluids of the host and frequently it constructs a

feeding tube for this purpose.

Exenterus abruptorius (Thnb.) (Ichneumonidae)

Large niunbers of Exenterus abruptorius (Thnb.) were imported by

Canada and released against the European spruce sawfly, and the

European pine sawfly. Large colonies were also made available for

release in the United States. A few specimens were recovered 7

years after liberation in New Jersey from Neodiprion sertifer, proving

definite establishment, but host populations in that area were reduced

almost to the vanishing point by an energetic spray program. Further

attempts at recovery have not been made. E. abruptorius is a very

important N. sertijer parasite in Europe.

Exenterus abruptorius has one generation a year. Adults are

active in early summer. The femfles embed their eggs under the

skin of prepupae or large larvae. Hatching does not take place

until the host larva has spun its cocoon. Larval development may be

retarded, but when it reaches maturity a, cocoon is spun within the

host cocoon and the winter is spent in this stage.

Exenterus spp. (Ichneumonidae)

Canadian entomologists successfully established _ four European

species of Exenterus: E. conjusus Kerrich, E. amietorius (Panz.),

È. veUicatus Cush., and E. tricolor Koman, against the European

spruce sawfly during the period of extensive parasite introduction

against that pest from 1934-49. Colonies were also made available

for release in the United States. Because of the sudden coUapse of

the inifestation, due principally to a virus disease, few attempts were
58 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

made to recover the parasites in this country, and the only one ever

reared in the united States was E. confusus from three cocoons collected in southern Vermont. Nevertheless, itseems proper to mention

all four species, because the fact that they are in New Brunswick

almost guarantees that eventually they will be found in the United

States, even though the sawfly remains at the low ebb at which it

has persisted for the last 20 years.

Exenterus confusus was the predominant species of Exenterus recovered in New Bnmswick, but it has been very rare in the extremely

low levels of host population prevailing since 1941. E. amictorius

was second to E. confusus in numbers, but it seems especially

dependent on host density and has not been recovered in recent

years. E. vellicatus is more capable than either E. confusus or E.

amictorius in maintaining itself at low host population levels. It has

been recovered in New Brunswick, Nova Scotia, and Newfoundland,

and has been consistently reared in small mnnbers from study plots

in New Brunswick. Only one specimen of E. tricolor was recovered.

The life histories of all four species are similar. Eggs are laid on

the integument of large larvae. Hatching normally occurs within

the host cocoon. The winter is usually spent as a last instar or

prepupa. In New Brunswick one and a partial second generation

develop annually with about 50 percent of the summer generation

remaining ia diapause untu the second year.

Exorista larvarum (L.) (Tachinidae)

Exorista larvarum (L.) is one of the important larval parasites of

the gypsy, brown-tail, and satin moths in Europe. It was extensively colonized in the United States from 1923 to 1932, but it was

not recovered untu 1940. In 1941 it was reared from larvae collected

in 11 different townships extending from Standish, Maine, to Waterbury, Conn. Since that time no extensive collections have been

made, and no figures are available regarding its importance. It is

multibrooded, and its value will depend to some extent on the abundance of alternate hosts. In Europe it has been reared from at least

45 different lepidopterous hosts, many of which are forest pests.

Exorista larvarum lays a lai^e white macrot3rpe egg on the integument of the host larva. The parasite maggot bores through the

skin and forms a respiratory funnel. When parasitizing the gypsy

moth, the full-grown larva emerges and drops to the groimd to form

its puparium. The puparium may be formed within the skin of the

host or the cocoon ofsome otherspecies. From one to several maggots

may develop in a single host larva. The winter is spent as an immature larva within an overwintering host.

Laricobius erichsoni Rosenh. (Derodontidae)

Laricobius erichsoni Rosenh. is a derodontid beetle. It preys upon

a number of adelgids in Europe, but its preferred prey is the balsam

woolly aphid. Canadian entomolorists have imported it since 1951,

and it has become generally established. They have also made

colonies available for release in New England, the Pacific Northwest,
PARASITES AND PREDATORS OF TOHEST INSECTS 59

and North Carolina. It is still too early to say whether or not it is

established in North Carolina, but it has been recovered at the other

liberation points. In 1959, 15,000 adults were released on the Penobscot Experimental Forest near Bangor, Maine, in an attempt to

use this predator for direct control on a limited area. This was a

cooperative project of the U.S. Forest Service (Division of Pest

Control and Northeastern Forest Experiment Station), U.S. Agricultural Research Service, and the Commonwealth Institute of Biological Control. It is too early to report on results.

Laricobius is an effective predator. At some liberation points in

New Brunswick, the spring generation of the aphid has been almost

completely destroyed and stem attacks have been terminated by the

beetles. The predator spreads very slowly, though, and is only

occasionally found in very light infestations. European workers do

not believe that Laricobius alone is capable of cleaning trees heavily

infested with the aphid.

Adult Laricobius overwinter in the duff. They attack all stages of

Chermes piceae except the first instar larvae. Eggs are laid in the

wax of the prey, and larvae feed on eggs and adult aphids. When

feeding is complete they drop to the ground to pupate. Newly

emerged beetles appear on the trees in early summer and feed some

before entering the soil to hibernate. Adult beetles live approximately one year.

Leucopis {Neoleucopis) obscura (Hal.)

(Chamaemyiidae)

Leucopis obscura (Hal.) is a chamaemyiid fly, native to Europe,

which was successfully introduced into Canada against the balsam

woolly aphid in 1933. It became established immediately and by

1947 was present throughout most of the infested area. It spread

naturally into Maine. In 1954 L. obscura was colonized and estabhshed in southern Vermont, and during 1955 and 1956 several more

colonies were released in Vermont, New Hampshire, and New York.

In 1958 and 1959 the fly was colonized in Washington and Oregon,

and in 1959 it completed at least a sununer generation in Oregon.

In 1960 it was released in North Carolina.

Leucopis is abxmdant in heavy balsam wooUy aphid stem mfestations. In 1958, for instance, an average of 15 Leucopis adults were

recovered per square foot of bark surface on thirteen 5-foot logs cut

from heavily infested trees near Bangor, Maine. Leucopis is dependent on high prey density, and it is seldom found except on rather

heavily infested stems. It faüs to prevent heavy oviposition because

it feeds mostly on adults that have laid many eggs. On the other

hand, it reduces the number of larvae produced in late stages ot

attack thus enabling some trees to recover, and it reduces the number

of crawlers dispersed from heavily infested trees Leucopis undergoes heavy winter mortaUty in Mame and New Brunswick. It wül

Be mteresting to know if it is more effective in warmer areas such

as Oregon and North CaroUna. It also attacks Pineus strobi (Htg.)

and P. pini Koch_in England.
60 tr.S. DEPT. OF AGRICtTLTXTRB HANDBOOK NO. 226

Leucopis completes two generations a year. The winter is spent

as a larva or puparium in crevices in the bark. Adults emerge in the

spring and deposit eggs singly in the "wool" of their prey. Larvae of

Letuiopis feed upon adelgid adults and larvae. A generation is completed in the spring, and adults emerge to attack the autumn generation of the aphid.

Mesoleius tenthredinis Morley (Iclmeumonídae)

Mesoleius tenthredinis Morley was first imported into Canada from

England in 1910-11 and liberated against the larch sawfly in Ontario

and Quebec. A small colony was also liberated on Grand Island

near Munising, Mich. In 1912-13 liberations were made in Manitoba.

This work was highly successful. According to A. R. Graham parasitization averaged 75 percent over a wide area in Manitoba in 1927

(Canad. Ent. v. 63, 1931).

About 1940 it became apparent that the effectiveness of Mesoleius

had decreased greatly in Manitoba and Saskatchewan. Apparently

the host developed a high immunity to the parasite in the general

region, because a very high proportion of the parasite's eggs became

encapsulated and failed to hatch. This immunity does not seem to

have developed as yet in British Columbia, the Maritime Provinces,

or Pennsylvania.

There is one generation of Mesoleius tenthredinis annually. The

winter is spent as a third or fomth instar larva within the cocooned

host larva. Feeding is resumed in the spring, and pupation takes

place within the host cocoon. Adults appear from late June to early

August and lay internal eggs in larch sawfly larvae. One parasite

develops in a host larva.

Meteorus versicolor (Wesm.) (Braconidae)

Meteorus versicolor (Wesm.) was one of the first brown-tail moth

parasites established in this country. It has become dispersed

throughout the whole of the infested area, but it never has parasitized

large numbers of this host. In fact, parasitism rarely exceeds 5

percent. In Europe M. versicolor also attacks satin moth larvae.

Attempts to establish it on the satin moth in New England have

failed, but it became quickly estabUshed on this host ia the Pacific

Northwest, and it is considered an important control factor there.

Parasitism of hibernating larvae up to 31 percent and of later instars

up to 17 percent were recorded at one point soon after establishment.

The possibiUty that Meteorus versicolor reared from the brown-taU

moth differs from M. versicolor reared from the satin moth cannot be

disregarded in spite of the fact that the two appear morphologically

identical.

Meteorus overwinters as a first instar larva in brown-tail moth

larvae. Development is completed in the spring, and large larvae

are in turn attacked. Undoubtedly other hosts are also parasitized

during the summer. In early fall hibernating brown-tail moth larvae

are again parasitized. The cocoon is spun in an unusual waj[. The

full-grown larva leaves the host and after crawling some little distance

along a twig or branch suspends itself by a long silken thread. The
PARASITES AND PREDATORS OF FOREST INSECTS 61

cocoon is spun in this position. It is thus fully exposed and consequently is often attacked by hyperparasites.

Monodontomerus aereus Walk. (Torymidae)

Monodontomerus aereus Walk, was introduced into New England in

1906 with brown-tail moth hibernation webs. It became established

readily and by 1910 was distributed over most of the infested area.

It was originally released as a primary parasite, and it does occasionally act as a primary internal parasite of lepidopterous pupae,

including the brown-tail and gypsy moths. It is a much more common hyperparasite, however, attacking both hymenopterous cocoons

and tachinid puparia, and it is often a serious hyperparasite of

such beneficial species as CompsUura.

There is usually only one generation a year. The winter is spent

as an adult, and brown-tail moth webs are a favorite hibernating site.

Ovipositing females usually place three or four eggs externally on the

host within the cocoon, puparium, or pupa, but when oviposition

occius in living lepidopterous pupae the eggs are deposited internally.

The females feed extensively on their hosts. A feeding tube extending

from the body of the host to the outer surface of the cocoon or puparium is formed with the ovipositor. Body fluids are sucked up through

this tube. Undoubtedly large numbers of hosts are kUled in this

way instead of being parasitized. The adults are long lived, often

Uving for 11 or more months.

Monodontomerus dentipes (Dalman) (Torymidae)

Monodontomerus dentipes (Dalman) is a European species, which

has been recorded from a number of hymenopterous and lepidopterous

hosts, and it frequently acts as a secondary parasite. It is, however,

an important primary parasite of several pine sawflies. It was already

parasitizing Diprion simüis (Htg.) when that pest was discovered in

the united States, and it is credited with being one of the most

effective checks upon that pest in New Jersey and Pennsylvania.

H. C. Goppel also reports that it is the most important biotic agent

controlling D. simüis in Wisconsin, although it was never released

there.

There are probably two generations of Monodontomerus dentipes

annually. The winter is spent as a prepupa inside the host cocoon.

Adults emerge in the spring, and there may be as much as a month's

difference between the emergence of the first and last adult from one

cocoon. The female oviposits through the host cocoon laying several

eggs externally upon the prepupa. The parasite larvae feed externaOy

completing development in 3 or 4 weeks. Four or five adults usually

emerge from one cocoon, but occasionally as many as twelve may

develop.

Ooencyrtus kuwanai (How.) (Encyrtidae)

OoencyHus kuwanai (How.), native to Japan, is an egg parasite of

the gypsy moth. Small numbers were introduced into this country m

1908 and 1909, and the species was very successfully reared m the
62 U.S. DEPT. OP AGRICULTURE HANDBOOK NO. 226

laboratory. By 1910 more than a million individuals were released.

The parasite became established almost immediately, and through

extensive colonization was dispersed throughout the infested area.

It is now considered an important parasite in the southern range of its

host. Parasitism of 40 to 45 percent of the eggs is frequently recorded

in Massachusetts and Connecticut.

In the laboratory Ooencyrtus kuwanai has been reared from the

eggs of several other Lepidoptera, but there are no records of it as an

egg parasite of other species in the field. It has been reared occasionally as a hyperparasite of Apanteles. It is poorly adapted to this host,

however, and there is no doubt that it is normally a primary egg

parasite.

Ooencyrtus spends the winter as an adult. A generation is completed

in the spring on overwintered gypsy moth eggs, and females are

sufficiently long lived to attack the newly laid eggs in July. At

least two generations are completed in the fall, and adult Ooencyrtus

are conspicuous on egg masses well into November in southeastern

Massachusetts and Connecticut. The female lays an egg within the

egg of the gypsy moth, and if the host larva is developed, as is usual,

the parasite egg is placed within the caterpillar. The host embryo is

entirely consumed except the hair and some hard chitinous parts.

Pupation takes place within the egg shell and the new adults cut their

way out.

Orgilus obscurator (Nees) (Ichneumonidae)

Orgüus obscurator (Nees) is one of the European pine shoot moth

parasites imported from Europe by Canada and the United States in

the thirties. It apparently had been accidentally introduced with

its host at an earlier date for it was reared from the shoot moth in

Rhode Island in 1925. There is little doubt, however, that importations were responsible for much of its rapid spread throughout

Connecticut, southern New York, and New Jersey. It is also well

established in Ontario and the Niagara region.

Orgilus seems to vary greatly in effectiveness. Percentage parasitism varied from 0 to 50 in 23 collections of host larvae overwintering

in 1946 and 1947 in Connecticut, southern New York, and New

Jersey. While it was present in 18 of the collections, average parasitization was only 7 percent. In western New York, near Chautauqua,

it was present in 7 of 11 collections, but average parasitization was

only 2 percent. In this country it has only been reared from the shoot

moth, but in Europe several other hosts are listed.

Orguus has a single generation each year. It overwinters as a

first instar larva within the hibernating host larva. When development is complete, it emerges from the host larva or pupa and spins a

thin white cocoon in the host larva's mine. The adidts oviposit

internally in the new generation of larvae.

Parasetigena silvestris R.-D. (Tachinidae)

Parasetigena silvestris R.-D. is one of the predominant parasites of

the gypsy moth in central Eimspe, Laige numbers were colonized

in the United States from 1924 to l§33, but definite establishment was
PARASITES AND PBBDATOES OF FOREST INSECTS 63

not confirmée! until 1937. By 1941 it had spread throughout most of

the infested region and at one point, where 500 gypsy moth larvae were

collected, 18 percent of them were parasitized by this species.

Parasetigena sUvestris has a single generation each year overwintering

as a puparium in the duff. Adult flies emerge in early spring. The

females lay large macrotype eggs on the cutide of the host larva, and

the parasite maggot bores through the skin forming a respiratory

funnel at the point of entry. The full-grown maggot drops to the

earth to form its puparium.

In Europe Parasetigena is also an important parasite of the nun

moth. It is occasionally reared from the brown-tail moth.

Phanomeris phyllotomae Mues. (Braconidae)

Phänomens phyüotomae Mues, was collected in Austria and liberated

against the birch leaf-mining sawfly in New England in the early

thirties. It became established at several liberation points but host

populations dropped drastically, and further checks on the effectiveness of the imported parasites were discontinued. In Austria P.phyllotomae parasitized a rathersmall proportion ofthePhyUotoma collected.

At one collection point parasitizations of 21 and 18 percent were

recorded in 1932 and 1933, but at most points it was rare.

Phanomeris has one generation a year. The winter is spent as a

larva within a cocoon spun in the host larval mine. Adults emerge

in early summer, and eggs are laid near the semiparalyzed host larva.

The parasite crawls to its host and feeds externally, completing feeding

in late summer or early fall.

Phobocampe disparts (Vier.) (Ichneumonidae)

Almost 300,000 cocoons of Phobocampe disparis were collected in a

gypsy moth infestation in Sicily in 1911 and 1912, and large colonies

of adults were liberated at several points in New England. The

species became established readily, but it has not dispersed widely

and, as a rule, only a small proportion of the larvae collected at any

one point is parasitized by this species. In fact, with the exception

of the early work, it has never again been recovered in large numbers

in Europe.

Phobocampe disparis has one generation a year. The winter is

spent as a cocoon on the forest floor. Adults attack flrst or second

instar larvae, and the full-grown parasites emerge from fourth instar

larvae and drop to the ground to spin their cocoons. The cocoons

are heavüy attacked by hyperparasites. There is also a considerable

mortality of parasite larvae within the host. These two factors

undoubtedly limit the potential effectiveness of the species tremendously. Phobocampe appears to favor the shaded part of infested

woodlands, and parasitism by this species is frequently much higher

in dense woodland than along the woods' edge.
64 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 228

Phytodietusfumiferanae Roh. (Ichneumonidae)

Phytodietus fumiferanae Roh. is an ichneumonid parasite of the

spruce budworm. It is distributed throughout western North America

from Arizona to British Columbia, but it does not occur in the East.

Rather high percentage parasitism by this species was recorded

locally m British Columbia between 1917 and 1920. From 1943 to

1947 Canadian entomologists collected large numbers of spruce budworm larvae in the same general area near Lillooet in order to procure

this species and others for liberation in the East. Similar work was

conducted between 1945 and 1949 by American and Canadian entomologists in Colorado and Oregon, and this work was further supplemented by laboratory rearing of Phytodietus in both countries. In

the United States almost all laboratory propagation was conducted

by the State of Maine. Colonizations were apparently unsuccessful,

for Phytodietus has not yet been reared from larvae collected in the

East, except a few, one year after releases were made at a poiut near

Belleville, Ont.

Phytodietus fumiferanae has only one generation a year. It overvvmters as a mature larva in a silken cocoon spun on the foliage.

Adults emerge in early summer, and the female lays an external egg

on the partially paralyzed host larva. The parasite larva feeds

externally, spinning its cocoon when fully grown.

Pimpla turíonelltie (L.) (Ichneumonidae)

Pimpla furionellae (L.) is a common pupal parasite of many Lepidoptera in Europe. In the early thirties it was reared in large numbers

from European pine shoot moth pupae in Austria. Adults sent to

this country from Austria arrived too late to attack shoot moth pupae

here, but it was successfully reared on pupae of Cacoecia cerasivorana

(Fitch) for general release. It has not been recovered in the United

States. Canadian entomologists have recovered a few specimens in

southern Ontario, but it does not appear to be well adapted to the

region.

Pimpla turionellae completed several generations a year in the

laboratory. The adult females lay an internal egg in the host pupa,

and the parasite larva feeds internally. Pupation takes place inside

the pupal shell. The winter is spent as an adult, and possibly at

times in an immature stage.

Scymnus (Pullus) impexus (Muís.) (Coccinellidae)

Scymnus (Pullus) impexus (Muls.) is a coccinellid beetle found from

central Europe to north Africa. It preys upon the balsam woolly

aphid, and Canadian entomologists have imported it each year since

1951 for release against that pest. They have also made colonies

available for release in southern New England, North Carolina, and

the Pacific Northwest. It has become established at most liberation

points in Canada and has spread short distances, but populations have

remained small.
PARASITES AND PREDATORS OF FOREST INSECTS 65

The life cycle of Scymnus impexus is well synchronized with that of

its host. Tùere is one generation a year. The winter is spent as an

egg, which is laid in bark crevices, or as an adult, which hibernates

ÍQ the bark particularly around old knots. Oviposition is practically

confined to the autumn. Adults usually live 12 to 13 months. Both

adults and larvae feed on aphid eggs, large larvae, and adults. Fullgrown larvae pupate on the bark. There seems to be heavy winter

mortality in Canada.

Temelucha interruptor (Grav.) (Ichueumonidae)

Temelucha interruptor (Grav.) [formerly called Cremastus interruptor Qrav.] is one of the most common parasites of the European

pine shoot moth in England and in Europe. Large numbers were

imported from England, Austria, and Holland in the thirties, and it

was well colonized in Canada and the United States. It became

established readily and recoveries were made in Connecticut, New

Jersey, and southern New York in 1937. Apparently, though, conditions are unsuitable for the parasite in this general region since it was

recovered in just 1 of 23 collections of host larvae overwintering

there in 1946 and 1947. The recovery was at Branford, Conn. Eight

of 100 larvae were parasitized.

Temelucha interruptor has been reared from several European hosts.

It completes one generation a year in Bhyacionia buoliana, overwintering as a first instar larva. Development is completed in the

spring, and full-grown larvae emerge from host larvae or pupae to

spin their delicate cocoons in the host larval mine. Adults are present

in Jime and July.

Tetrastichus hrevistigma Cahan (Eulophidae)

Teirastichus hrevistigma Gahan is a native species that heavily

parasitizes the pupae of the imported elm leaf beetle in New England.

It was first discovered in 1932 near Boston, Mass. Percent parasitization increases toward the end of summer. In 1934 and 1935, by

late August, it averaged 76 and 47 percent, respectively, at five collection points. It was colonized in California in 1934 and became

established immediately. It is now also abundant in the infested

areas in that State.

Tetrastichus hrevistigma overwinters as a full-grown larva in the

dead host pupa, and an average of 12 larvae complete development

in one pupa. The parasite passes through three to four partial generations a year, the second generation adults emerging about the time

that beetle pupation is at its peak.

Tetrastichus turionum (Htg.) (Eulophidae)

Tetrastichus turionum (Htg.) is a European parasite of the European

pine shoot moth which was collected in large numbers in Austria in

the early thirties, and colonized throughout the infested area in New

England, southern New York, and New Jersey. It was recovered in

Massachusetts, New Jersey, and on Long Island. Canadian entomologists have recovered it from southern Ontario, where they had
66 U.S. DEPT. OF AGKICULTUBB HANDBOOK NO. 226

liberated it. In spite of early recoveries the parasite is apparently

poorly adapted to conditions in the United States, for it has been

absent or very rare in numerous collections made to determine its

effectiveness.

Tetrasfichus turionum is a gregarious pupal parasite that completes

one generation annually. The full-grown larvae overwinter in a

tightly packed mass within the pupal shell of the host. Pupation

occurs in the spring, and adult females emerge about the time the host

larvae are pupating. Females oviposit an average of about 20 eggs

internally in each host pupa.

Townsendiellomyia nidicola (Tns.) (Tachinidae)

Tovmsendiellomyia nidicola (Tns.) [=Alsomyia nidicola (Tns.)] was

one of the first brown-tail moth parasites established in this countiy.

It spread rapidly throughout the infested areas, and now it frequently

parasitizes an average of 17 percent of the hibernating larvae. In

some collections parasitism runs as high as 40 percent. The browntail moth is the only known host in this countiy.

Tovmsendiellomyia nidicola has one generation a year. The winter

is spent as a first instar maggot in the overwintering host larva.

Development is completed in the fuU-grown larva, and the puparium

is formed in the dead host larva's skin. First instar maggots are

fully developed in the eggs that the female deposits on the venter of the

small host larvae.
INDEX OF HOST SPECIES

Adelges piceae, 2

Alsophila pometaria, 24

Anticarsia sp., 24

aphid, balsam woolly, 2, 4, 54, 58, 59, 64

Asterolecanium—

hambusae, 2

variolosum, 2

balsam fir sawfly, 23

balsam wooly aphid, 2, 4, 54, 58, 59, 64

bamboo scale, 2

elm leaf, 16, 18, 56, 65

southern pine, 9

birch leaf-mining sawfly, 20, 22, 53, 63

brown-tail moth, 27, 29, 52, 54, 66, 57,

58, 60, 61, 63, 66

budworm, spruce, 3, 6, 64

Cacoecia cerasivorana, 64

Carpocapsa pomonella, 24

casebearer, larch, 9, 10, 53

caterpillar—

forest tent, 20

great basin tent, 20

range, 20, 21

Chemes piceae, 2, 4, 49, 55, 59

Choristoneura—

fumiferana, 3, 6

Tnuriana, 3

Cnidocampa flavescens, 3, 8

Coleophora laHcella, 9, 10, 53

Colzas pkilodice eurytheme, 25

Datana integerrima, 24

Dendroctonus frontalia, 9

Diprion—

frutetorum, 9, 55

hercyniae, 9, 12, 50, 55, 56

polyiomum, 16

similis, 16, 61

Douglas-fir tussock moth, 17, 21

elm leaf beetle, 16, 18, 56, 65

elm leaf miner, 53

elm scale, European, 17, 53

Epilachna varivestris, 24

European elm scale, 17, 53

European pine sawfly, 9, 26, 28

European pine shoot moth, 32, 38, 54,

62, 64, 65

European spruce sawfly, 9, 12, 55, 56,

57

Exoteleia dodecella, 54

Fenusa ulmi, 53

forest tent caterpillar, 20

Galerucella xanthomelaena, 16, 18

golden oak scale, 2

GossypaHa spuria, 17

great basin tent caterpillar, 20

gypsy moth, 27, 30, 33, 50, 51, 52, 54

56, 58, 61, 62, 63

Harrisina hrillians, 25

HeTnerocampa—

leucosHgma, 17, 19

pseudotsugata, 17, 21

Hemileuca oHviae, 20, 21

Heterarthrus nemoratus, 20, 22, 53

Hyphantria cunea, 24, 25

introduced pine sawfly, 16

jack-pine sawfly, Swaine's, 27

Laphygma frugiperda, 25

larch casebearer, 9, 10, 53

larch sawfly, 32, 60

Malacosoma, 20

disstria, 20

fragilis, 20

pluviale, 20

Miresa niiens, 37

moth—

brown-tail, 27, 29, 52, 54, 56, 57, 58,

60, 61, 63, 66

Douglas-fir tussock, 17, 21

European pine shoot, 32, 38, 54, 62,

64,65

gypsy, 27, 30, 33, 51, 52, 54, 56, 58,

61, 62, 63

Nantucket pine tip, 51

nun, 63

oriental, 3, 8, 52

pine tip, 37

satin, 37, 44, 54, 57, 58

wMte-marked tussock, 17, 19

Nantucket pine tip moth, 51

Neodiprion—

ahikis, 23, 55

fvlviceps, 23

lecontei, 23, 26, 55, 56

nanulus, 55

pini-rigidae, 55

pratti paradoxiciis, 26, 50, 55

praiti pratti, 26, 55

sertifer, 26, 27, 28, 50, 55, 57

swainei, 27

nettle grub, 37

nun moth, 63

Nygmia phaeorrhoea, 27, 29, 31

oriental moth, 3, 8, 52

Phryganidia californica, 24

Phyllotoma (see Heterarthrus).

Pikonema—

alaskensis, 56

dimmockii, 56

pine sawfly—

European, 9, 26, 28

introduced, 16

red-headed, 23

67
68 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

pine shoot moth, European, 32, 38, 54,

62, 64, 65

pine tip moth, 37, 51

Pineus—

pint, 59

Portheiria dispar, 27, 30, 33

Pristiphora erichsonii, 32

range caterpillar, 20, 21

red-headed pine eaurfly, 23

Rhopobota naevana, 24

Rhyacionia—

buoliana, 32, 38, 54, 65

frustrana, 51

frustraría hushneîîi, 37, 51

frustraría neomexicana, 37, 51

satin moth, 37, 44, 54, 67, 58, 60

sawfly—

balsam ñr, 23

birch leaf-mining, 20, 22, 63, 63

European pine, 9, 26, 28

European spruce, 9, 12, 65, 56, 57

sawfly—continued

introduced pine, 16

larch, 32, 60

red-headed pine, 23

Swaine's jack-pine, 27

scale—

bamboo, 2

European elm, 17, 53

golden oak, 2

Setora nitens, 37

southern pine beetle, 9

spruce budworm, 3, 6, 64

spruce sawfly, European, 9,12, 55, 56, 57

Stüpnoiia scuicis, 37, 44, 57

Swaine's jack-pine sawfly, 27

tent caterpillar—

forest, 20

great basin, 20

tussock moth—

Douglas-fir, 17, 21

white-marked, 17, 19

white-marked tussock moth, 17, 19
INDEX OF INSECT ENEMIES

Actia nudibasis, 38

Adalia—

luteopicta, 4

tetraspilota, 4

Agathis pumüus, 9, 10, 47

Agria affinis, 6

Alsomyia nidicola (see Townsendiellomyia nidicola).

Anastatus disparis, 30, 32, 33, 47

Apanteles, 57, 62

lacteicolor, 27, 29, 48

liparidis, 30, 33

melanoscelus, 30, 32, 33, 48

porthetriae, 30

solitarius, 17, 19, 30, 43, 44, 49

Aphidecta obliterata, 4, 49

Aphidoletea thompsoni, 3, 4, 49

Apíesís—

basizonia, 12, 16, 26, 27, 28, 50

subguttata, 13

Ballia eucharis, 4

Blepharipoda scutellata, 30, 32, 33, 50

Blondelia—

nigripes, 30, 33

piniariae, 29, 33

Brachymeria—

míermedía, 30, 33

o&scwraíor, 30

Calliephialtes comstocki (see Ezeriates

comsiockii).

Calosoma—

chínense, 30

inquisitor, 30

reticulatum, 30

sycophanta, 17, 20, 21, 23, 24, 27, 30,

32, 33, 34, 44, 51

Campoplex, 38

borealis, 38

frustranae, 37, 42, 51

multidnctus, 38

nvutabilis, 38

Tufifemur, 38

Carabws—

arcensts, 30

aryents (see Carabus arcensts).

auratus, 29, 30, 32, 52

glabratus, 30

nemoralis, 30, 52

violaceus, 30

Carcelia—

^nawo, 44

laxifrons, 29, 30, 52

rutila, 29

separata, 30, 34

Ceromasia auricaudata, 6

Chaetexorista javana, 3, 8, 37, 52

ChiUtcorus kuwanae, 4

Chrysis shanghaiensis, Z, 8

Ckrysocharis, 22

laricinellae, 9, 10, 20, 22, 53

Coccophagus insidiator, 17, 53

Compsilura condnnata, 6, 17, 19, 20, 21,

23, 25, 27, 29, 30, 32, 34, 43, 48, 54

Copidosoma geniculatum, 38, 54

Cremasíiís—

epagoges (see Temelucka epagoges).

interruptor (see Temelucha interruptor).

Cremifania nigrocellulaia, 3, 4, 54

Crossocosmia—

flavosciUellata, 30

sericariae, 30

Dahlhominus fuscipennia, 9, 13, 15, 16,

23, 26, 27, 28, 32, 55

Delomerista—

diprionis (see D. japónica),

japónica, 13, 28

DicladoceriLS westwoodi, 11

Diomus pumilio, 5

2)rmo bohémica, 16, 26, 31, 36, 56

EphiaUes ruficoUis, 39

Eremochila ruficoUis (see EphiaUes

ruficoUis).

Ernestia sp., 35

Erynniopsis rondani, 16, 56

Eupteromalus nidulans, 29, 43, 45, 57

Eurytoma pint, 42

Exenterus, 14

abruptorius, 13, 26, 27, 28, 57

amictorius, 14, 16, 28, 57

claHpennis, 14, 16

confusus, 57

pmi, 14

/ricoior, 14, 16, 57

veUicatus, 16, 57

Exeristes comstockii, 42

Exochomus—

¿¿¿uroius, 4

uropygialis, 4

Exorista—

larvarum, 27, 29, 30, 31, 32, 35, 43,

45,58

segregata, 30

Glypta varipes, 42

Habrocytus thyridopterigis, 42

Habrolepis dalmani, 2

HalticheUa rhyacioniae, 42

Harmonía hreiti, 5

Horogenes—

laricinellae, 11

nana, 11

Hyssopus—

rhyacioniae, 42

thymus, 42

Ischnojoppa luteator, 14, 28
70 U.S. DEPT. OF AGRICULTURE HANDBOOK NO. 226

lioplectis 4:-cingulatus, 6, 14

Lamachus, 14

eques, 14, 28

marginatuSf 14

Laricohius erichsoni, 3, 5, 58

Lehia scapularia, 18

Lets dimidiaia, 5

Leucopis (Neoleucopis), 2, 5

obscura, 2, 3, 6, 59

Lixophaga—

mediocrisj 42

plúmbea, 42

Lophyropectus Ivieator (see Ischnojoppa

luieator).

Lydelta, 14

nigripes (see Blondelia nigripes).

piniariae (see Blondelia pimariae).

Lypha dubia, 39

Macrocentrus sp., 39

Masicera sylvaiica, 30

Mesoleius tenthredinis, 32, 60

ilfeieorws—

japonicus, 30

pulchricornis, 31

versicolor, 27, 29, 43, 45, 60

Microterys sp., 17

Monodontomerus—

aereus, 29, 32, 61

dentipes, 15, 16, 61

Neoleucopis obscura, 2, 3, 5, 59

Olesicampe ratzeburgi, 15

Ooencyrtus kuwanai, 17, 19, 21, 31, 32,

35, 47, 61

Orgilus obscurator, 37, 39, 62

Paies pavida, 31, 46

Parasetigena silvestris, 31, 32, 35, 62

Phanomeris phyllotomae, 20, 22, 63

Phobocampe, 35

disparis, 31, 32, 35, 63

Phorocera kamata, 15

Phytodietus fumiferanae, 3, 6, 64

Pimpla—

examinator, 31

instigator, 31

turionellae^ 17, 20, 21, 37, 40, 65

Pristomerus vulneraior, 40

Procrustes coriacßus, 31

Prosturmia hohemica, 16, 66

Pullus impexvs, 3, 5, 64

Rogaa unicolor^ 46

5ca7n6tts foliae, 22

Scymnus—

imp&cits, 3, 5, 64

pumiliOf 5

Spathimeigenia spinigera, 15

Sturmia, 15

inconspicua, 26, 31, 36, 56

scuteüata (see Blepharipoda scutellaia).

Synharmonia conglóbala, 5

Tachina—

japónica, 31

mo^nicornis, 31

7bchim<¿8, 31

TocAinomyia similis, 46

Tcíenomus pkalaenarum, 31

TVrraeiiicÄo—

epagoges, 42

irUerruptor, 37, 40, 54

Tetrastickus, 16, 19

hrevisiigma, 16, 18, 65

gcUerucae, 16, 18

¿«rionûm, 37, 41, 65

xardhomelaenae (see T. galerucae).

Thanasimvx formicarius, 9

ToionseíwiieZíomí/xa nidicola, 27, 29, 66

TVanosema pedella, 22

TWcfcofframma, 31

evane5C£7i5, 41

TVícAomasíAus cyanifrons, 17

Xylodrepa quadripunctaia 31

Zenillia libairix, 31, 36, 46

U.S. GOVERHHEHT PRINTIK6 OFriCBi IS«!
Y!»ii .,.