i,'

^

-M

United States

Department of

Agriculture

Combined Forest Pest

Research and

Development Program

Agriculture Handbook

No. 540

¡573^96^

Gypsy Moth

Handbook

Selected

Parasites

and

Hyperparasites

of the Gypsy

Moth,

with Keys to

Adults

and Immatures
In 1974 the US. Department of

Agriculture initiated the Combined Forest

Pest Research and Development Program,

an interagency effort that concentrated on

the Douglas-fir tussock moth in the West,

on the southern pine beetle in the South,

and on the gypsy moth in the Northeast.

The work reported in this publication was

funded in whole or in part by the program.

This manual is one in a series on the gypsy

moth.

usofc.«»***'

Möo-»««'^***

^^^>rx^
Contents

Preface 4

Introduction 5

Keys to Adults of Selected Parasites and

Hyperparasites of the Gypsy Moth 7

Key to families 7

Key to Tachinidae 8

Key to Ichneumon idae 11

Key to Braconidae 13

Key to Chalcididae 14

Key to Encyrtidae, Eupelmidae, Scelionidae, and Torymidae 15

Key to Selected Inunatures 16

Tables of Biological Information for Selected Parasites of

the Gypsy Mothft^^^^^ ^ ^ ., 20

Table 1. Species known*tô bh-ésX^^h^d in or native to North America 20

Table 2. Species not known to be estabfi^ed in ^North.. America 24

Notes on Species and Names not Included in Keys orvTal)le^ ^ 30

Glossary ^-.-'^Í' •^•^.í'-K-'^' ' " ' • 33

References '**"-^VOi ;,»f '"' '"•''-'"/ 39

Figures " ' "' '• ''á';:.f," „», 45

Acknowledgments 59
Selected Parasites and

Hyperparasites of the Gypsy

Moth,^ with Keys to Adults

and Immatures

by

Edward E. Siinons,^ Richard C.

Reardon,^ and Mark Ticehurst^

Preface

In recent years there has been

renewed interest in biological

control of destructive forest insects,

including the gypsy moth. Studies of

gypsy moth parasites have been

hampered by lack of keys to

parasitic wasps and, until recently,

to parasitic flies. This handbook

includes keys to adults, puparia, and

cocoons of selected gypsy moth

parasites found in, imported to, or

likely to be imported to North

America. The keys are

supplemented by illustrations and

by a technical glossary. Also

included are tables summarizing

important biological data on all

parasites treated in the keys.

Additional notes are made

concerning why certain species

were not included, as well as

synonyms used in the North

American literature.

^ Lepidoptera: Lymantriidae. Lymantria

dispar (L.) is preferred to the often used

name Porthetria dispar (Ferguson 1978fl).

2 Entomologists, Pennsylvania Department

of Environmental Resources, Middletown,

Pa.

^ Entomologist, Forest Insect and Disease

Management, Forest Service, Hamden,

Conn.
Introduction

Parasite importation for the

biological control of Lymantria

dispar (L.) began in 1905 and

continued through 1914. Further

importations were made from 1922

to 1933, and from 1%3 to the

present. More than 40 species have

been introduced; 9 are currently

established in the Northeastern

United States. The gypsy moth

importation and release programs

have been reviewed by Howard and

Fiske (1911), Burgess and Grossman

(1929), Dowden (1%2), Hoy (1976),

Clausen (1978), and Reardon (1978).

Lymantria dispar is distributed in

the western Palearctic region from

southern Sweden and Russia south

of 58° N latitude to North Africa,

Syria, and northern Iran; and in the

eastern Palearctic region from

Siberia south of 55° N latitude to

Japan, China, northern India, and

northern Afghanistan (Anon. 1953).

In the Nearctic region, L. dispar is

distributed at present in New

England except northern New

Hampshire, northern Maine, and

northeastern Vermont; in

southeastern Quebec; in New

Jersey; in eastern and central New

York and Pennsylvania; in

northeastern and northcentral

Maryland; and in southcentral

Michigan (Baker 1972, Bryce 1978).

The typical L. dispar of the western

Palearctic and Nearctic regions is

replaced by Lymantria dispar

japónica (Motschulsky) in Japan

(Ferguson 1978fc). Lymantria

obfuscata (Walker) is distributed in

India (Sabrosky and Reardon 1976).

The parasites in this handbook

include flies and wasps known to

parasitize the gypsy moth and

include native, established, and

exotic species recently released and

other selected exotic species.

In this handbook are keys to

families and species of adults and

immatures that are designed to aid

in the identification of parasites

recovered from L. dispar They

cannot be satisfactorily used to

identify suspected gypsy moth

parasites whose hosts are unknown.

Selected hyperparasites are included

in the keys if they are likely to be

recovered from field-collected gypsy

moth caterpillars and pupae.

Hyperparasites that attack primary

parasites after they leave the gypsy

moth are not included.

Hyperparasites are identified in the

keys but are not discussed further in

the handbook. Schewyrew (1912)

has been followed in regard to the

two species of Theronia and

Monodontomerus aereus Walker as

hyperparasites.
The key to immature stages is

limited to those forms that are

external to the host. Some closely

related species could not be

separated because of morphological

similarities. Despite these

limitations, the immature key should

be useful when adults are in poor

condition, fail to emerge, or when

immediate identifications are

required.

The keys are intended to be useful

to workers with some training in

insect morphology and taxonomy.

Illustrations and a glossary are

provided to illustrate and explain

technical terms; both should enable

the reader to use the keys without

references to other publications. In

the use of morphological

terminology, recent authors have

been followed in the taxonomy of

particular groups included here:

Tachinidae from Sabrosky and

Reardon (1976); and Hymenoptera

from Townes (1%9) and Joseph et

al. (1973).

Information on biology, distribution,

and relative importance is presented

in tabular form for all parasites

included in the keys. Biological data

are based on personal observation

and on world literature. The biology

and relative importance of the

parasites may vary with location,

host density, and other variables.

This information is not intended to

be complete but to serve as a guide

for further reference.

keys and tables. Thompson (1946)

lists 186 species of wasps and flies

reared from the gypsy moth. Other

sources of parasite records are

Shenefelt (1972), Telenga (1955),

Moriey and Rait-Smith (1933),

Muesebeck et al. (1951), Krombein

(1958), and Krombein and Burks

(1%7). Not all catalogers have

critically reviewed host and

distributional records; some data,

therefore, are inaccurate (Nixon

1974). The criteria mentioned earlier

eliminate most of these species from

consideration, although it may

superficially appear that some

species not included do meet the

criteria mentioned earlier. Reasons

for not including certain species are

given in a separate section, which

also includes comments on

synonyms of species treated here

that were used in North American

literature. Only wasp synonyms are

discussed in this section, because

synonyms of gypsy moth fly

parasites are well covered by

Sabrosky and Reardon (1976).

There are numerous parasites that

have been recorded from the gypsy

moth besides those treated in the
Keys to Adults of Selected

Parasites and Hyperparasites

of the Gypsy Moth

Key to Families

1. With one pair of wings; hind wings reduced to haltères. Diptera

(flies) 2

— With two pair of wings, or so tiny as to be difficult to determine the

number of wings. Hymenoptera (wasps) 3

2. Postscutellum strongly convexly developed Tachinidae

— Postscutellum not strongly convexly developed Other Diptera^

3. Front wings with both longitudinal and cross veins enclosing at least

a few cells (figs. 34 and 43) 4

— Front wings with venation reduced to an apparent single longitudinal

vein having a short branching vein near the end, and without closed

cells (figs. 53 and 54) 5

4. Two recurrent veins in front wing (fig. 34) Ichneumonidae

— One recurrent vein in front wing (figs. 43 and 44) Braconidae

5. Body length^ usually at least 4 mm long; hind femora swollen and

dentate on lower margin (fig. 52) Chalcididae

— Smaller, usually less than 4 mm long; hind femora normal size or

possibly with a single tooth on lower margin 6

6. Pronotum in side view triangular and extending to the tegula (fig.

61) Scelionidae

— Pronotum in side view quadrate and not reaching the tegula (fig. 62) 7

7. Hind coxa considerably larger than front coxa; mesopleura with a

distinct groove for the reception of the femora (fig. 55); body length

over 2 mm Torymidae

— Hind coxa little if any larger than front coxa; mesopleura usually

large and convex, at least without a distinct groove (fig. 56); body

length between 1 and 2 mm 8

8. The submarginal vein long and extending to the base of the stigmal

vein, without a marginal vein (fig. 57); the mesonotum without notauli and evenly convex (fig. 58) Encyrtidae

— The submarginal vein shorter, with a marginal vein (fig. 53); mesonotum with notauli and either convex (fig. 59) or flattened or slightly

concave with a median triangular elevation anteriorly (fig.

60) Eupelmidae

^ To separate scavengers see Sabrosky and

Reardon(1976).

•' Body length is measured from the

anterior part of the head exclusive of the

antennae to the apex of the abdomen

exclusive of the ovipositor.
Key to Tachinidae

1. Two sternopleural bristles (fig. 20)^ 2

— 3 or 4 sternopleural bristles (figs. 17, 18, and 19) 4

2. Intermediate abdominal segments without median discal bristles;

cheeks unusually narrow (fig. 6); legs predominantly yellow, only

tarsi black Carcelia amplexa (Coquillett)

— Intermediate abdominal segments with median discal bristles (fig. 4);

cheeks not as narrow (figs. 7, 8, and 9) 3

3. Mesonotum, viewed from behind, with 4 narrow black stripes (fig.

2); each humérus with 3 strong bristles arranged in a triangle (fig.

15) Eusisyropa virilis (Aldrich and Webber)

— Mesonotum, viewed from behind, with 3 stripes, the median a fiision

of usual pair of submedian stripes, and twice breadth of a lateral

stripe (fig. 3); each humérus with 5 bristles (fig. 16)

Nemorilla pyste (Walker)

4. Intermediate abdominal segments, each with pair of strong and erect

median discal bristles; abdomen of female with spined ventral keel

(fig. 26) 5

— Intermediate abdominal segments without median discal bristles; abdomen of female without spined ventral keel 6

5. Eyes appearing bare, their hairs minute and sparse (fig. 10); 3 pairs

of posterior dorsocentral bristles Blondelia nigripes (Fallen)

— Eyes densely long haired; 4 pairs of posterior dorsocentral

bristles Compsilura concinnata (Meigen)

6. Four sternopleural bristles (fig. 18) 7

— Three sternopleural bristles 10

7. Facial ridges bristled on lower V2 to ^U\ wing with 2-3 bristles on

node at base of R4+5 on upperside of wing; underside of fourth abdominal segment of male of normal appearance, the hairs well

spaced 8

— Facial ridges almost bare, only a few weak bristles and hairs above

vibrissae; wing with only one bristle on node at base of R4+5; underside of fourth abdominal segment of male with a pair of well-defined,

dense fascicles of hairs (figs. 24 and 25) 9

8. Paired median marginal bristles present on abdominal segments 1+2

and 3 (fig. 4), the pair on 1+2 normally weak

Lespesia frenchii (Williston)

— Median marginals absent on abdominal segments 1 +2 and 3

^ Rarely a weak, hairlike, third

sternopleural bristle in Eusisyropa virilis.

If in doubt, E. virilis has 3 strong humeral

bristles arranged in a definite triangle (fig.

15).
Lespesia aleîiae (Riley)

9. Hair patches, on underside of fourth abdominal segment, in males

large, each obviously more than half as long as segment; the area

between patch and anterior margin of segment bare of hairs (fig.

24) Palexorista inconspicua (Meigen)

— Hair patches in males small, each V2 length of segment (fig. 25); with

at most a row of ordinary hairs between patch and anterior margin

of segment, or with only scattered hairs, or none at all

Palexorista disparis Sabrosky

10. Prealar bristle long and strong, obviously longer than adjacent

intra-alar bristle and as long as posterior notopleural (fig. 13); 1 pair

of reclinate orbital bristles present; eyes bare; abdomen with median

marginal bristles on segments 1+2 and 3, those on 3 especially

strong Blepharipa pratensis (Meigen)

— Prealar bristle relatively short, usually clearly less than adjacent intra-alar bristle and posterior notopleural bristle (fig. 14); 2

pairs of reclinate orbital bristles present (fig. 4) 11

11. Eyes distinctly haired (figs. 11 and 12) 12

— Eyes appearing bare, hairs minute and sparse (Fig. 10) 14

12. Back of head with a partial row of black hairs behind postocular

cilia Parasetigena silvestris (Robineau-Desvoidy)

— Back of head with only whitish hairs behind postocular cilia 13

13. Facial ridges with strong, well-spaced bristles on lower % to ^U\

second antennal segment bright reddish yellow in both sexes

Spoggosia claripennis (Macquart)

— Facial ridges not as strongly bristled, bristles weaker, closer together,

and decumbent, on lower half or less; second antennal segment

black except narrowly at apex Exorista segregata (Rondani)

14. Two median lateral bristles present on each side of scutellum (fig.

21) Tachinomyia spp.

— Only 1 median lateral bristle on each side of scutellum (fig. 22) .. .15

15. Dorsum of abdomen almost entirely gray tomentose, without conspicuous shining black crossbands, and with a more or less distinct

median white line, or partial line Exorista rossica Mesnil

— Dorsum of abdomen with broad, shining black crossbands posteriorly on segments 3-5, and usually with distinct black median line

or narrow stripe, often less evident in females 16
16. Males^ with median forceps (fused cerci) characteristically elongate,

narrow, compressed (fig. 28); parafrontals anterioriy, especially in

males, with fine, pale hairs, appearing almost bare under low magnification; cheek in both sexes relatively narrow, as seen in profile

only Vs height of eye Exorista japónica (Townsend)

— Males with median forceps (fused cerci) broadened and flattened,

distally suddenly narrowed to acute apex (fig. 29); parafrontals, especially in males, with longer black hairs that are much more distinct; cheek in both sexes usually wider, about V4 height of

eye Exorista larvarum (L.)

Exorista mella (Walker)"

' Females of the 3 following species are

difficult to separate.

^ The relationship of palearctic E.

larvarum, extensively introduced into

North America, and the native E. mella

has not been satisfactorily studied.

10
Key to Ichneumonidae^

1. Areolet large, with one of its sides usually as long as or longer than

upper abscissa of postnervulus (fig. 30); first tergite with a large

glymma, its spiracle near or a little behind the middle; male clasper

ending in a narrow rod about half the length of first segment of the

hind tarsus. Hyperparasite Mesochorinae^"

— Areolet usually smaller and narrower (fig. 31), all of its sides plainly

shorter than upper abscissa of postnervulus; otherwise not entirely

agreeing with above 2

2. First abdominal segment somewhat petiolate, usually at least twice

as long as widest part, and at least twice as wide at apex as near

base viewed from above, spiracle situated behind the midlength of

the tergite (fig. 32) 3

— First abdominal segment quadrate, length less than twice the width,

apex but little wider than base viewed from above (fig. 33); spiracle

situated near or before the midlength of the tergite (fig. 34) 6

3. Second abdominal tergite with a gastrocoelus

Lymantrichneumon disparis (Poda)

— Second abdominal tergite without a gastrocoelus 4

4. Cross section of first abdominal segment near its basal third depressed oval in shape; suture separating first abdominal sternite from its

tergite above the midheight on the petiolar part of segment (fig. 35);

propodeum viewed from above narrowed toward apex, dorsal surface without distinct carinae except at base; inner margin of eyes

strongly indented near base of antennae (fig. 36) Casinaria spp.

— Cross section of first abdominal segment near its basal third quadrate or triangular in shape; suture separating first abdominal sternite

from its tergite below the midheight on the petiolar part of segment

(fig. 39); propodeum viewed from above quadrate in overall shape,

not appreciably narrowed toward apex; median longitudinal carinae

extending beyond basal area; inner margin of eyes not strongly indented 5

5. Petiole of first abdominal segment with a conspicuous lateral pit or

glymma (fig. 37); abdominal tergites entirely black; nervulus of front

wing forming a wider angle of at least 70° with the discoideus (fig.

38) Hyposoter tricoloripes (Viereck)

— Petiole without a conspicuous lateral pit (fig. 39); second abdominal

" Sections of this key were adapted from

an unpublished key by Robert Carlson.

'" For a recent taxonomic study of this

subfamily see Dasch (1971).

11
tergite with apex brown; nervulus strongly sloping to make an angle

of about 55° with the discoideus (fig. 31)

Phobocampe disparis (Viereck)

6. Thorax and abdomen mostly yellow or light brown; mesopleural suture with a weak angulation near the middle (fig. 34); tarsal claws

enlarged, each with an enlarged bristle with a spatulate tip (fig. 40) ..

Theronia spp. 7

— Thorax and abdomen mostly black; mesopleural suture straight; tarsal claws not enlarged and not with a spatulate bristle 8

7. Head mostly black, conspicuously darker than most of thorax; hind

femur without a ventral ridge. Hyperparasite .. Theronia hilaris (Say)

— Head yellowish, concolorous with most of thorax; hind femur with a

ventral ridge (fig. 34). Hyperparasite

Theronia atalantae fulvescens (Cresson)

8. Apices of abdominal tergites whitish; base of most hind tarsal segments white, contrasting with dark apices; inner margin of eye

strongly concave slightly above level of insertion of antennae (fig.

41) Itoplectis conquisitor (Say)

— Apices of abdominal tergites dark; hind tarsal segments entirely

dark; inner margin of eye usually only weakly concave (more strongly concave in males of C. turionellae and C disparis) (fig. 42)

Coccygomimus spp. 9

9. Hind tibia with a whitish area medially; dark basally and apically . 10

— Hind tibia entirely black, fuscous, or reddish yellow 11

10. Hind coxa and disc of scutellum reddish

Coccygomimus turionellae moraguesi (Schmiedeknecht)

— Hind coxa and disc of scutellum black

Coccygomimus turionellae turionellae (Linnaeus)

11. Hind coxa reddish yellow Coccygomimus pedalis (Cresson)

— Hind coxa black 12

12. Hind tibia reddish yellow Coccygomimus instigator (Fabricius)

— Hind tibia fuscous or black Coccygomimus disparis (Viereck)

12
Key to Braconidae

1. Front wing with 2 cubital cells (second intercubital vein absent) (fig.

43); body usually not over 3 mm long Apanteles spp. 4

— Front wing with 3 cubital cells (second intercubital vein present although sometimes indistinct) (fig. 44); body usually at least 4 mm

long 2

2. First abdominal tergum widened at apex, at least 3 times as wide as

apex as at base (fig. 45); clypeus with apical margin transverse and

touching upper edge of mandibles Meteoms pulchricornis (Wesmael)

— First abdominal tergum with apex less than twice as wide as base

(fig. 46); clypeus semicircularly arched at apical margin, forming a

circular opening with mandibles (fig. 47) Rogas spp. 3

3. Body mostly medium to light brown .... Rogas indiscretas Reardon

— Body mostly dark brown to dull black, dorsal surfaces black except

disc of second abdominal segment and sometimes apex of first abdominal segment and parts of mesonotum and scutellum brown

Rogas lymantriae Watanabe

4. Apex of first abdominal tergum narrower than base (fig. 48); propodeum mostly smooth and shining, irregularly punctate but not rugose 5

— Apex of first abdominal tergum as wide or wider than base (fig. 49);

propodeum rugose 6

5. First abdominal tergum evenly tapered at apex (fig. 48)

Apanteles porthetriae Muesebeck

— First abdominal tergum abruptly narrowed at apex (fig. 50)

Apanteles liparidis (Bouché)

6. Second abdominal tergum about Vio longer than third tergum; third

tergum usually coarsely punctate to apex; basal antennal segments

of female bright yellow Apanteles ocneriae Ivanov

— Second abdominal tergum subequal to Vio shorter than third tergum;

third tergum usually distinctly smoother and less punctate on apical

half; basal antennal segments of female black

Apanteles melanoscelus (Ratzeburg)

13
Key to Chalcididae^^ (genus

Brachymeria)

1. Complete frontal (preorbital) carinae present in both sexes^^ (fig. 51);

females without a median (inner ventral) tooth on hind coxae; apical

half of scutellum with a vaguely defined, median, longitudinal depression; space between lateral ocellus and compound eye shagreened (covered with a closely set roughness). Hyperparasite

B. compsilurae (Crawford)

— Frontal (preorbital) carinae absent in females (or vaguely indicated

and short in males); if present in females then hind coxa with a

median (inner ventral) tooth 2

2. Hind coxae of female with a median (inner ventral) tooth (fig. 52) . .3

— Hind coxae without an inner tooth or protuberance both in male and

female 4

3. Hind tibia yellow with a blackish ventral carina from base to tip;

interspaces between the punctures on the thorax smooth

B. intermedia (Nees)

— Hind tibia mostly yellowish with the base and ventral portion blackish; interspaces between the puncture on the thorax finely recticulate B. lasus (Walker)

4. Scutellum emarginate; preorbital and postorbital carinae distinct;

hind tibia yellowish at base, apex and dorsal portion, and ventral

middle portion black. Hyperparasite B. fiskei (Crawford)

— Scutellum weakly emarginate or entire or rounded; preorbital carina

faint or absent; postorbital carina present; hind tibia mainly yellow

with the base alone black B. euploeae (Westwood)

" Taken, in part, from Joseph et al. (1973)

although we have followed the opinion of

Boucek (1973) in treating B. euploeae and

B. lasus as distinct species.

^^ B. fiskei also has preorbital carinae

present in both sexes (see couplet 4).

14
Key to Encyrtidae, Eupelmidae,

Scelionidae, and Torymidae

1. Pronotum in profile more or less triangular and extending to the

tegulae (fig. 61); trochanter 1-jointed 2

.., Anastatus disparis Ruschka

— Pronotum in profile more or less squarish and not extending back to

the tegulae (fig. 62); trochanter 2-jointed 3

2.^^ Abdomen aside with blunt lateral edge; frons smooth and polished;

females with 11-segmented antennae Telenomus spp. ^^

— Abdomen aside with sharp lateral edge; frons roughly sculptured; females with 12-segmented antennae Gryon spp.^^

3. Mesopleura large, entire; flat, without femoral groove in the female

and usually in the male (fig. 56); spur of middle tibia usually very

large and stout 4

— Mesopleura rarely large, with an oblique femoral groove or impression (fig. 55); spur of middle tibia normal, not enlarged 5

4. Submarginal vein long and extending to the base of stigmal vein;

without a marginal vein (fig. 57); 11-segmented antennae

Ooencyrtus kuvanae (Howard)

— Submarginal vein shorter and with a marginal vein (fig. 53); 12- or

13-segmented antennae Anastatus kashmirensis^^ Mathur

Anastatus disparis Ruschka

5. Hind coxae more or less triangular in cross-section, sharply ridged

above; hind femora slightly swollen with a single ventral tooth.

Hyperparasite Monodontomerus aereus Walker

— Hind coxae cylindrical, long; hind femora greatly swollen and with

many teeth beneath Brachymeria spp. (see key to Chalcididae)

i^Taken, in part, from Masner (1958).

14 The male oí Telenomus lymantriae

Kozlov is unknown and the female has

first abdominal segment and legs, including

coxae, yellow; body black.

i^Two species oí Gryon, howardi

Mokrzecki and Ogloblin and lymantriae

Masner, have been infrequently recovered

from the gypsy moth, and their exact

relationship has not been satisfactorily

studied. The genus Gryon is included here

to avoid confusion with Telenomus.

i^The females are similar morphologically,

while the males are distinct: \nA.

kashmirensis, the hind tibial basal area is

tan and the apical areas dusky, and '\nA.

disparis, the hind tibia is dark except at

the extreme base. Final determination

pending examination of topotypical

material of kashmirensis-disparis-japonicus

(Gordh 1976).

15
Key to Selected Immatures

The species and groups treated in this key are limited to those in the resting

stage that are more or less visible. In some genera and tribes, characteristics

could not be found to separate species, as was the situation for Rogas spp.

and the Exoristini. Parasetigena silvestris (R.-D.) is the only common gypsy

moth parasite in the Exoristini. Differences in biology should help separate

P. silvestris from any Exorista spp. and Spoggosia claripennis that may be

reared from gypsy moth, because the latter two have at least two generations a year, whereas P. silvestris has only one. The only species of Careelia

treated in this key is amplexa (Coquillett), because it is the only one reared

from gypsy moth that is known to be in the Nearctic region. The tachinid

section of the key was modified from Sabrosky and Reardon (1976).

16
1. Pupa covered at least partially with silken strands (i.e. cocoon),

color white, yellow, light brown, gray or black; or within mummified

skin of host caterpillar; wasp cocoons 2

— Pupa covered with skin of last stage of the larva, without silken

strands, color reddish brown to black; fly puparia 9

2. Cocoon within mummified skin of host caterpillar Rogas spp.

— Cocoon external to host although host remains may adhere to it .. .3

3. Cocoons 4-5 mm long; thickly covered with silken strands; color

white or pale yellow Apanteles spp. 4

— Cocoons usually at least 5 mm long; covering parchmentlike, with

only a few silken strands, or at least sections of darker parchmentlike covering visible beneath silken strands; at least partly brown,

gray, or black ^

4. Cocoons occurring in groups (a cluster of at least 4 or 5)

Apanteles ocneriae Ivanov 2inà Apanteles liparidis (Bouché)

— Cocoons occurring singly ^

5. Cocoons pale yellow Apanteles melanoscelus (Ratzeburg)

— Cocoons pure white Apanteles porthetriae Muesebeck

6. Cocoon a uniform brown color; free from host, and suspended from

substrate at end of a silken strand; 2 mm in diameter and 5-6 mm

long; ends acutely rounded (fig. 73) Meteoms pulchricornis Wesmael

— Cocoon with mottling, banding or both; if color neariy uniform then

it is dark gray; free or adhering to host; usuaUy at least 3 mm in

diameter; ends bluntly rounded 7

7. Cocoon tightly adhering to ventral side of caterpillar remains (fig.

74) Hyposoter tricoloripes (Viereck)

— Cocoon free from host 8

8. Cocoon length at least twice width; with irregular dark bands or

mottling that contrasts with white or yellow coloration elsewhere . ..

Casinaria spp.

— Cocoon stouter, length only about IV2 times diameter; with one

broad, gray median band (sometimes indistinct) and dark gray apices

(fig. 75) Phobocampe disparis (Viereck)

9. Puparium densely beset with spinules, or fine spinelike hairs, which

form a furiike covering; spiracular plates strongly projecting, height

one half or more the diameter of plates; the surface of spiracular

plates very uneven with the slits at the apex of 3 ridges (fig.

54) Palexorista spp. 10

— Puparium chiefly bare, any spinules confined to definite rows or

narrow segmental bands; spiracular plates usually flush with surface

or at least if projecting with the surface relatively smooth 11

17
10. Spiracular slits straight or only slightly curved; the distance between

plates about Vs the diameter of plates; no fold or integument surrounding the area of spiracular plates

Palexorista inconspicua (Meigen)

— Spiracular slits wavy; the distance between plates about equal to

diameter of plates; 1 or 2 folds of integument surrounding the area of

spiracular plates Palexorista disparis Sabrosky

11. Spiracular slits strongly sinuous, with distinct loops (fig. 65) 12

— Spiracular slits straight or only gently curved (fig. 66) or wavy or

zigzag (fig. 67), but not widely looped 14

12. Subspiracular protuberance large, prominent, and with triangular,

ridgelike extension dorsally between spiracular plates (fig. 65)

Blepharipa pratensis (Meigen)

— Subspiracular protuberance, if present, without triangular extension

between plates Lespesia spp. 13

13. Subspiracular protuberance moderately strong (fig. 68)

Lespesia aletiae (Riley)

— Subspiracular protuberance small and weak (fig. 69)

Lespesia frenchii (Williston)

14. Spiracular slits distinctly wavy or zigzag in outline (figs. 67 and 70) 15

— Slits straight or gently curved (fig. 66) 16

15. Strong subspiracular protuberance present (fig. 67)

Carcelia amplexa (Coquillett)

— Protuberance absent (fig. 70) Eusisyropa virilis (Aldrich and Webber)

16. Posterior end of puparium narrowed, subconical, the spiracular

plates distinctly projecting and slightly divergent (fig. 71)

Nemorilla pyste (Walker)

— Posterior end of puparium not subconical, the spiracular plates little

if any elevated above surrounding surface (figs. 63 and 66) 17

17. Posterior end of puparium in profile somewhat depressed above

apex and more or less conspicuously bulging between anus and subspiracular protuberance (fig. 63); spiracular plates well above apex

of puparium, on the depressed and slanting surface

Tribe Exoristini (Exorista spp., Parasetigena, Spoggosia,

Tachinomyia)

— Posterior end of puparium in profile broadly and evenly rounded, the

spiracular plates almost vertical to long axis of puparium (fig. 66) 18

18. Usually with deep groove between spiracular plates, plates comparatively small, with small subspiracular protuberance close to them

and usually projecting beyond them (fig. 72)

Blondelia nigripes (Fallen)

18
Space between plates flat or finely wrinkled, not with deep groove;

plates comparatively larger, and subspiracular protuberance less projecting and not as close to plates, the 3 appearing less crowded together (fig. 66) Compsilura concinnata (Meigen)

19
Tables of Biological

Information for Selected

Parasites of the Gypsy Moth

Table 1.—Species Known to Be

Established in or Native to North

America

Parasite

Stage

attacked

Host

preference Voitinism

Overwintering

stage

Braconidae

Apanteles

melanoscelus

(Ratzeburg)

larva oligophagous bivoltine immature in

cocoon

Ichneumonidae

Coccygomimus

pedalis

(Cresson)

pupa polyphagous multivoltine prepupa (?) in host

Itoplectis

conquisitor

(Say)

pupa polyphagous multivoltine prepupa (?) in host

Phobocampe

disparis

(Viereck)

larva oligophagous univoltine adult in

cocoon

Encyrtidae

Ooencyrtus

kuvanae

(Howard)

egg oligophagous multivoltine female adult in

litter

Eupelmidae

Anastatus

disparis

Ruschka

egg oligophagous univoltine

(occasionally

bivoltine)

immature in host

Chalcididae

Brachymeria

intermedia

(Nees)

pupa polyphagous multivoltine female adult in

litter

Tachinidae

Blepharipa

pratensis

(Meigen)

larva oligophagous univoltine pupa in litter

Carcelia

amplexa

(Coquillett)

larva polyphagous uni-bivoltine pupa in litter

Compsilura

concinnata

(Meigen)

larva polyphagous multivoltine immature in host

20
Distribution and Relative

importance References

western Palearctic and

Northeastern United States,

parasitism commonly 0-22%

Grossman 1922, Burgess and

Grossman 1929, Ticehurst et al.

1978

Nearctic, parasitism occasional Townes and Townes 1%0

Nearctic, parasitism occasional Townes and Townes 1%0

western Palearctic and

Northeastern United States,

parasitism commonly 0-30%

Burgess and Grossman 1929,

Ticehurst et al. 1978, Muesebeck

and Parker 1933

Palearctic and Northeastern United

States, parasitism commonly 10-

50%

Burgess and Grossman 1929,

Dowden 1%1, Grossman 1925

Palearctic and some areas in

Northeastern United States,

parasitism commonly 0-40%

Burgess and Grossman 1929,

Parker 1933, Grossman 1925

Neotropic, Palearctic, and

Northeastern United States,

parasitism commonly 0-67%

Joseph, Narendran and Joy 1973,

Dowden 1935, Ticehurst et al.

1978, Burks l%Oa, Krombein and

Burks 1%7

Palearctic and Northeastern United

States, parasitism commonly 0-54%

Sabrosky and Reardon 1976,

Ticehurst et al. 1978

Nearctic, parasitism occasional Sabrosky and Reardon 1976

Palearctic and Nearctic, parasitism

commonly 0-54%

Sabrosky and Reardon 1976,

Ticehurst et al. 1978

21
Parasite

Stage

attacked

Host

preference Voltinism

Overwintering

stage

Eusisyropa

virilis (Aldrich

and Webber)

larva polyphagous bivoltine immature in host

Exorista

larvarum (L.)

larva polyphagous multivoltine immature in host

Exorista mella

(Walker)

larva polyphagous multivoltine immature in host

Lespesia aletiae

(Rüey)

larva polyphagous multivoltine immature in host

Lespesia

frenchii

(WiUiston)

larva polyphagous multivoltine immature in host

Nemorilla pyste

(Walker)

larva polyphagous multivoltine immature in host

Tachinidae

Parasetigena

silvestris

(RobineauDesvoidy)

larva oligophagous univoltine pupa in litter

Spoggosia

claripennis

(Macquart)

larva polyphagous multivoltine pupa in litter

Tachinomyia

spp.

larva polyphagous univoltine pupa in litter

22
Distribution and Relative

importance

Nearctic, parasitism occasional

References

Sabrosky and Reardon 1976

Palearctic, parasitism common;

United States, parasitism rare

Nearctic, parasitism occasional

Sabrosky and Reardon 1976

Sabrosky and Reardon 1976

Nearctic, parasitism occasional Sabrosky and Reardon 1976

Nearctic, parasitism occasional Sabrosky and Reardon 1976

Nearctic, parasitism occasional Sabrosky and Reardon 1976

Palearctic and Northeastern United

States, parasitism commonly 0-68%

Sabrosky and Reardon 1976,

Ticehurst et al. 1978

Nearctic, parasitism occasional Sabrosky and Reardon 1976

Nearctic, parasitism occasional Sabrosky and Reardon 1976

23
Table 2.—Species Not Known to

Be Established in North America

Parasite

Stage

attacked

Host

preference Voltinism

Overwintering

stage

Braconidae

Apanteles

liparidis

(Bouché)

larva polyphagous multivoltine immature in host

Apanteles

porthetriae

Muesebeck

larva oligophagous multivoltine immature in host

Apanteles

ocneriae

Ivanov

larva monophagous univoltine(?) immature in

cocoon(?)

Meteorus

pulchricornis

Wesmael

larva polyphagous multivoltine

Rogas

indiscretas

Reardon

larva oligophagous bivoltine(?) immature in

mummified host

Rogas

lymantriae

Watanabe

larva monophagous bivoltine(?) immature in

mummified host(?)

Ichneumonidae

Casinaria

spp.

larva polyphagous multivoltine prepupa (?) in

cocoon

Coccygomimus

disparis

(Viereck)

pupa polyphagous multivoltine prepupa(?) in host

Coccygomimus

instigator

Fab.

pupa polyphagous multivoltine prepupa(?) in host

Coccygomimus

turionellae

turionellae

(L.)

pupa polyphagous multivoltine prepupa(?) in host

24
Distribution and Relative

importance References

Palearctic, parasitism common Burgess and Grossman 1929

western Palearctic, parasitism

common

Burgess and Grossman 1929,

Muesebeck 1928

western Palearctic, parasitism

occasional

Telenga 1955, Vasic 1973

western Palearctic, parasitism

occasional

Burgess and Grossman 1929

eastern Palearctic (India),

parasitism occasional

Reardon et al. 1973

eastern Palearctic (Japan),

parasitism occasional

Palearctic, parasitism occasional Townes, Momoi and Townes 1%5,

Fuester 1978

eastern Palearctic, parasitism

occasional

Townes, Momoi and Townes 1%5

Palearctic, parasitism occasional Townes, Momoi and Townes 1%5

Palearctic, oriental parasitism

occasional; Nearctic (Ganada)

Townes, Momoi and Townes 1%5,

Walkley 1958, Oehlke 1%7

25
Parasite

Stage

attacked

Host

preference Voltinism

Overwintering

stage

Coccygomimus

turionellae

moraguesi

(Schmiedeknecht)

pupa polyphagous multivoltine prepupa(?) in host

Hyposoter

tricoloripes

(Viereck)

larva polyphagous multivoltine prepupa(?) in

cocoon

Lymantrichneumon disparis

(Poda)

female adult in

decaying logs,

stumps, etc.

Eupelmidae

Anastatus

kashmirensis

Mathur

egg polyphagous multivoltine larva in host

Chalcididae

Brachymeria

lasus

(Walker)

pupa polyphagous multivoltine

Brachymeria

euploeae

(Westwood)

pupa polyphagous multivoltine

Sceiionidae

Telenomus

spp.

egg polyphagous multivoltine

Gryon spp. egg polyphagous multivoltine

Tachinidae

Blondelia

nigripes

(FaUen)

larva polyphagous multivoltine immature in host

Exorista

japónica

(Townsend)

larva polyphagous multivoltine immature in host

26
Distribution and Relative

importance References

western Palearctic (Morocco) Oehlke 1%7

western Palearctic, parasitism

occasional

Fuester 1978

Palearctic, parasitism occasional Howard and Fiske 1911

eastern Palearctic (India),

parasitism occasional

Oriental and eastern Palearctic Joseph et al. 1973

eastern Palearctic (India) Joseph et al. 1973

Palearctic, parasitism occasional

Palearctic, parasitism occasional

Kozlov 1%7

Masner 1958

Palearctic, parasitism occasional Sabrosky and Reardon 1976

eastern Palearctic, parasitism

occasional

Sabrosky and Reardon 1976

27
OverParasite

Stage

attacked

Host

preference Voltinísm

wintering

stage

Exorista larva polyphagous multivoltine immature in host

rossica

Mesnîl

Exorista larva polyphagous multivoltine immature in host

segregata

Rondani

Palexorista larva polyphagous multivoltine immature in host

disparis

Sabrosky

Palexorista larva polyphagous multivoltine immature in host

inconspicua

(Meigen)

28
Distribution and Relative

importance References

Palearctic, parasitism common Sabrosky and Reardon 1976

western Palearctic, parasitism

common

Sabrosicy and Reardon 1976

eastern Palearctic (northern India),

parasitism common

Sabrosky and Reardon 1976

Palearctic, parasitism occasional Sabrosky and Reardon 1976

29
Notes on Species and Names

Not Included in Keys and

Tables

Amistcitus hifasciatus Fonscolmbe

of Howard and Fiske (1911),

Grossman (1925), Muesebeck and

Dohanian (1927); misidentification

oíAnastatus disparis Ruschka. See

Burks (1967) and Pfeck (1963).

A nastatusjaponicus Ashmead.

Specialists are uncertain if this

eastern Palearctic species is the

same as/i. disparis Ruschka. If

they are the same sptcitsj'aponicus

is the preferred name because it has

priority (Gordh 1976).

Apantelesfulvipes (Haliday) of

Howard and Fiske (1911);

misidentification oíA. liparidis

(Bouché). See Burgess and

Grossman (1929).

Apanteles inclusus Ratzeburg. This

widely distributed Palearctic species

is a parasite oíEuproctis

chrysorrhoea (L.) (browntail moth),

Lymantria monacha (L.) (nun

moth), and Euproctis similis

(Fuessly) (goldtail moth). Records of

it irom Lymantria dispar (L.)

(Shenefeit 1972) were not

confirmed.

Apanteles lacteicolor Viereck. This

western Palearctic species was

introduced early and became

established on the browntail moth in

New England (Burgess and

Grossman 1929). Muesebeck (1918)

found that parasitism by the first

generation adults upon small gypsy

moth caterpillars could be

considerable when in the vicinity of

a browntail infestation. However,

the choice of gypsy moth seems to

be one only of necessity to find an

alternate host to complete the

summer generation. The wasp is still

limited in distribution to the

relatively small area of New

England colonized by the browntail

moth. For workers collecting in that

area the adult can readily be

separated from otherApanteles

reared from gypsy moth caterpillars

by the presence of the areola on the

propodeum (Muesebeck 1920).

Apanteles ruidus Wilkinson.

Gotypes in the United States

National Museum were reared in

India from Pyrausta machaeralis.

Shenefeit (1972) includes no species

oíLymantria in the host list. The

laboratory colony at the

Gonnecticut Agriculture Experiment

Station, New Haven, Gonnecticut,

called '^Apanteles ruidus^^

originating in India is not ruidus of

Wilkinson (1928) according to

Muesebeck (Weseloh 1978). The

taxonomic status of the New Haven

colony (and other colonies obtained

from the same population) is

uncertain, but is close to A.

melanoscelus (Ratzeburg).

Apanteles solitarius (Ratzeburg) of

Howard and Fiske (1911), Parker

(1935), and Dowden( 1962);

synonym of^4. melanoscelus

(Ratzeburg). See Nixon (1974).

Apanteles vitripennis (Gurtis) of

Grossman and Webber (1924),

Burgess (1926); misidentification of

A. porthetriae Muesebeck

(Muesebeck 1978).

Brachymeria euploeae (Westwood).

At present, colonies of a

30
Brachymeria from India are being

maintained at Middietown, Pa., and

New Haven, Conn. The species was

originally identified as B. euploeae

(Westwood). However, individuals

key to B. lasus in Joseph,

Narendran, and Joy (1973). Lab

studies indicate that it readily

parasitizes Exorista japónica

puparia as well as L. dispar pupae

when given a choice of the two,

whereas B. lasus, in colony at

Middietown and originating in

Japan, prefers L. dispar pupae,

although it will parasitize E.

japónica in the absence ofL. dispar

(Fusco 1978). Therefore, the

taxonomic status of this

Brachymeria population from India,

although close to lasus, is still

uncertain.

Brachymeria obscurata (Walker) of

Burgess and Grossman (1929), Hoy

(1976), and Thompson (1954);

synonym of ^. lasus (Walker). See

Joseph, Narendran, and Joy (1973).

Campoplex validus (Cresson) of

Krombein and Burks (1967);

transferred to Sinophorus by

Townes(1969).

Chalcisflavipes Panzer of Crawford

(1910), Howard and Fiske (1911),

Crossman and Webber (1924);

misidentification ofBrachymeria

intermedia (Nees). See Mariatt

(1928) and Burks (1960b).

Chalcis obscurata Walker of

Howard and Fiske (1911), and

Crossman and Webber (1924);

synonym ofBrachymeria lasus

(Walker). See Joseph, Narendran,

and Joy (1973).

Coccygomimus tenuicorn is

(Cresson). Riley and Howard (1894)

report occasional parasitism of the

gypsy moth. However, this was

probably a misidentification of C.

pedalis (Cresson) or Itoplectis

conquisitor (Say) (Carison 1978).

Ephialtes (Itoplectis) conquisitor

(Say) of Cushman (1920b);

transferred to Itoplectis by Townes

(1940).

Ephialtes pedalis (Cresson) of

Cushman (1920b); transferred to

Coccygomimus by Townes (1945).

Ephialtes temnopleuris Cushman

(1920b); synonym ofItoplectis

conquisitor (Say). See Townes

(1940).

Ephialtes tenuicorn is (Cresson) of

Cushman (1920b); transferred to

Coccygomimus by Townes and

Townes (1960).

Eulimneria valida (Cresson) of

Thompson (1946); transferred to

Sinophorus by Townes, Townes, and

Gupta (1961).

Glyptapantelesfulvipes (Haliday) of

Howard (1905); misidentification of

Apanteles liparidis (Bouché). See

Burgess and Crossman (1929).

Hyposoter disparis Viereck (1911)

of Burgess and Grossman (1929),

and Muesebeck and Parker (1933);

transferred to Phobocampe by

Townes (1945). The name //.

disparis is synonymized by Carlson

(1979).

31
Ichneumon disparis Poda of

Howard and Fiske (1911), and

Thompson (1946); transferred to

Lymantrichneumon by Heinrich

(Í968).

Limnerium disparis Viereck of

Howard and Fiske (1911);

transferred to Phobocampe by

Townes (1945).

Meteomsjapónicas Ashmead of

Burgess and Grossman (1929); and

Hoy (1976); synonym of M.

pulchricornis. See Muesebeck

(1978).

Meteoms versicolor Wesmael. This

species from the western Palearctic

region was introduced early and

became established on the browntail

moth (Burgess and Grossman 1929).

Gypsy moth caterpillars are only

occasionally attacked by this species

(Muesebeck 1918).

Ooencyrtus kuwanai (Howard) of

Muesebeck et al. (1951), and

Krombein and Burks (1967).

Incorrect emendation of O.

kuvanae. See Goulson (1978).

Pimpla conqidsitor (Say) of Howard

and Fiske (1911); transferred to

lîoplecùs by Johnson and Hammar

(1912).

Pimpla disparis Viereck (1911) of

Howard and Fiske (1911);

transferred to Coccygomimus by

Townes and Townes (I960).

Pimpla examinator Fab. of Howard

and Fiske (1911) and Dowden

(1962); synonym oíCoccygomimus

turionellae (L.). See Townes,

Momoi, and Townes (1965).

Pimpla instigator (Fab.) of Morley

and Rait-Smith (1933), Thompson

(1946), and Dowden (1962);

transferred to Coccygomimus by

Townes and Townes (1960).

Pimpla pedalis Gresson of Fernald

(1896), Howard and Fiske (1911),

and Townes (1940); transferred to

Coccygomimus by Townes (1945).

Pimpla porthetriae Viereck (1911)

and of Howard and Fiske (1911);

synonym oíCoccygomimus disparis

(Viereck). See Townes, Momoi, and

Townes (1965).

Pimpla tenuicornis Gresson of

Fernald (1896); transferred to

Coccygomimus by Townes (1960).

Protichneumon disparis (Poda) of

Morley and Rait-Smith (1933);

transferred to Lymantrichneumon

by Heinrich (1968).

Schedius kuvanae Howard (1910)

and of Howard and Fiske (1911),

Grossman (1925), Muesebeck and

Dohanian (1927), Burgess and

Grossman (1929); transferred to

Ooencyrtus by Mariatt (1929).

Sinophorus validus (Gresson). This

species has been reared from gypsy

moth, but only rarely (Garison

1973).

Theronia fulvescens (Gresson) of

Howard and Fiske (1911), Gushman

(1920a), Brimley (1938); status

changed to Theronia atalantae

32
Glossary

fulvescens (Cresson) by Viereck

(1917).

Theronia melanocephala (Brülle) of

Fernald (1896), Viereck (1917),

Cushman (1920úí), Cushman (1928),

Brirtiley (1938); synonym of

Thewnia hilaris (Say). See

Cushman and Gahan (1921) and

Townes (1940).

This glossary provides an

explanation of technical terms used

in the keys. Both singular and plural

(pi.) forms of a word are given if the

plural is unlike English plurals

ending in -s. Synonymous terms are

denoted by (==).

abdomen.—^The posterior, or third

section of the insect body. In the

parasitic wasps the section is

narrowly joined to the posterior of

the thorax. In hymenoptera the

apparent first section is the

morphological second segment, the

morphological first segment having

been transferred to the thorax as the

propodeum. In some species the

abdomen is abruptly enlarged so

that the narrow "waist'' is not

apparent. Abdominal segments are

numbered from front to rear with

the apparent first segment

designated number one. In the

tachinids the abdomen is apparently

composed of four segments. The

apparent first is a composite with

the second and referred to here as

segment 1 +2, the following ones as

3,4, and 5. Segments 3 and 4

(apparent 2 and 3) are referred to as

the intermediate segments.

abscissa.—A section of wing vein

from one interruption to the next.

antenna, pi. antennae.—Paired

appendages on the head above the

face. In the tachinids they are three

segmented, with the apical segment

longer than the basal two. In the

wasps they are many segmented,

sometimes elbowed, and usually

attached about the midheight of the

head, but sometimes are attached

33
only a short distance above the

mouth.

apical.—At the end, tip, or

outermost part.

areolet.—A small cell (second

cubital cell) near the center of the

front wing of some ichneumonid

wasps (fig. 30).

carina, pi. carinae.—A ridge or keel

(fig. 51).

cereus, pi. cerci (anal forceps).—

The median lobes of the male

genitalia (figs. 28 and 29).

cheek (=gena).—Side of head

below the eye (fig. 7).

clasper.—^The lateral, outside piece

of the male genitalia.

clypeus.—A sclerite on the lower

part of the anterior of the head

between the face and the labrum,

usually separated from the face by a

groove.

cocoon.—A papery or silken

structure enclosing the pupa.

cubitus.—^The third longitudinal vein

in the apical half of the front wing

which in the wasps typically has its

base near the base of the stigma and

its apex a little posterior to the apex

of the wing. The basal abscissa is

often absent (fig. 43).

discal bristles.—^The bristles on the

central area of the dorsum of

abdominal segments (fig. 4).

decumbent.—Referring to bristles

that are bent over or angled so as to

form about a 45° angle or less with

the surface to which they are

attached.

discoideus.—In the front wing of

wasps the vein running as the

apparent continuation of the médius

and curving to near the posterior

margin of the wing (fig. 31).

dorsum.—^The back or top side of

an insect.

emarginate.—Notched.

eye.—^The large compound eye on

each side of the head composed of

many individual elements (fig. 4).

coxa. pi. co.xae.-

leg segment.

-The first (basal)

Cubital cells.—The cells of the front

wing of wasps lying between the

radius and cubitus. There are

typically three with the first lying

behind the stigma and the third at

the apex of the wing. They are

separated by the first and second

intercubital cross veins (fig. 44).

34
face.—Anterior of the head between

the eyes and between the base of

the antennae and the clypeus (fig.

41).

femoral groove.—A dorso-ventral

concavity on the mesopleuron

where the mesofemur normally

rests when appressed against the

thorax (fig. 55).

femur, pi. femora.—^The third leg

segment, the more basal of the two

longest leg segments (fig. 52). In

many species the second segment is

very small, making the femur the

apparent second segment.

first intercubitus.—^The first (and

sometimes only) cross vein

connecting the radius and cubitus

(fig. 44).

first recurrent vein.—In the front

wing of wasps the cross vein

connecting the cubitus with the

discoideus (fig. 44).

gastrocoelus.—A concave

impression on each side near the

base of the second tergite of wasps.

glymma.—A groove or pit on some

wasps in the side of the first tergite,

between its spiracle and base (fig.

37).

head.—^The first main body section,

having mouth parts, antennae, and

eyes.

humérus, p\. humeri.—The shoulder

or convex anterior corner area of

the mesonotum (fig. 5).

intra-alar bristles.—A row of

bristles on each side of mesonotum

lateral to a dorsocentral row (fig. 4).

keel.—Spined ridge on underside of

abdomen, formed by the

compressed ventral parts of

segments; shaped like the keel of a

ship (fig. 26).

mandible.—One of the paired

mouth-part structures in wasps.

They are hinged to the lower part of

the head a short distance below the

eyes (fig. 41).

marginal bristles.—^The bristles on

the hind margins of segments,

usually as a pair of median

marginals on abdominal segments

1+2 and 3 (fig. 4).

marginal vein.—In the front wings

of some wasps a longitudinal vein

running along the anterior edge from

the apex of the submarginal vein to

the base of the stigmal vein (fig. 54).

médius.—A longitudinal vein in the

basal part of the front wing about

midway between the front and hind

margins (fig. 31).

35
mesonoîum.—The dorsal part of the

relatively large middle section of the

thorax, and partially bordered

laterally by the base of the front

wings in tachinids and most wasps.

mesopleuml suture.—A vertical or

somewhat oblique groove near and

more or less parallel to the posterior

edge of the mesopleurum. It runs

from the base of the middle coxa to

the base of the front wing (fig. 34).

mesopleuron.—The relatively large

middle section of the side of the

thorax.

metanotum.—^The dorsal area of the

last thoracic segment. In wasps

narrow and indistinct and lying

between the postscutellum and the

much larger propodeum.

nervulus.—The cross vein which

connects the discoideus and

submedius in the front wing of

wasps (fig. 38).

notaulusy pi. notauli.—One of a pair

of grooves of the mesonotum of

some wasps separating the median

area from the lateral areas (fig. 59).

ocellusy pi. ocelli—^The small simple

eye located at the top of the head.

There are typically three with the

center more anterior than the lateral

two (fig. 41).

parafrontai—^The area of the head

above the base of the antennae

between the median stripe and the

eye (fig. 5).

pectinate.—Referring to a structure

having a row of close set spines, like

the teeth of a comb.

petiole.—In wasps the anterior part

of the first abdominal segment

extending back to the spiracle (fig.

37).

posterior dorsocentral bristles.—

Two rows of bristles on the posterior

area of the mesonotum. They are

the second row of bristles lateral to

the median line (fig. 4).

postnervulus.—In the front wing of

wasps the cross vein which

connects the posterior end of the

first recurrent vein to the posterior

longitudinal vein (brachius) (fig. 30).

postocular cilia.—Slender even

hairs in a row immediately behind

each eye, usually with tips curved

forward over the eye (fig. 4).

postscutellum.—A transverse area

immediately behind or beneath the

scutellum; in tachinids,

conspicuously convex and bulging

(fig. 5).

pre-alar bristle.—^The anterior

postsutural bristle of the row at the

edge of the mesonotum immediately

above the wing base (figs. 4, 13, and

14).

propodeum.—The apparent last

dorsal section of the thorax in wasps

(fig. 32). It is the morphological first

36
segment of the abdomen which has

transferred to the thorax. It extends

anterodorsally almost to the

postscutellum, with the metanotum

being narrow and indistinct.

pupa.—^The inactive (''resting')

stage in those insects (such as true

flies and wasps) that transform from

a larva to an adult of different

appearance.

radius.—^The second longitudinal

vein in the apical half of the wing.

Its base typically joins the stigma

and its apex terminates at the wing

margin anterior to the apex of the

wing (fig. 43).

reclínate orbital bristles.—Bristles

inclined backward which are located

between the anterior ocellus and the

eye (fig. 4).

sclerite.—A hardened body wall

plate bounded by sutures or

membranous areas.

scutellum.—A median,

subtriangular or shield-shaped area

projecting at the rear of the main

area of the mesonotum (fig. 5).

second intercubitus.—The more

posterior cross vein connecting the

radius and cubitus, if the cubital cell

is interrupted by two veins (fig. 44);

absent in some wasp groups

(Apanteles spp.).

second recurrent vein.—In the front

wing of ichneumonids the cross vein

connecting the cubitus to the

subdiscoideus (fig. 34).

spatulate.—Narrowed basally and

broad and flattened apically.

spiracle.—An external opening of

the trachial system. In adult

tachinidae there is one on each side

of the propleura, and on the

metapleura. In adult parasitic wasps

there is one at the lateral edges of

the propodeum and the first seven

abdominal segments (fig. 32).

spiracularplates {=stigmatal

plates).—In mature larvae and

puparia of Diptera, the hardened

and usually black plates at the

posterior bearing the spiracular slits,

usually flush with surface of

puparium or only slightly elevated,

sometimes strongly projecting (fig.

65).

37
spiracular slits.—Narrow slitlike

spiracles occurring on the spiracular

plates. They occur in sets of three in

most tachinids (fig. 73).

Stemite.—Hardened body wall plate

on the ventral side of the body

especially of an abdominal segment.

sternopleural bristles.—The bristles

on the sternopleuron, or central

area of side of thorax, above the

coxa of middle leg (fig. 17).

stigma.—^A thickening of the wing

membrane near the middle of the

front margin of the front wing. It is

usually darker than the membrane

(fig. 30).

stigmal vein.—Tht short vein

extending posteriorly from the front

margin in the front wings of some

wasps (fig. 57).

subdiscoideus.—A longitudinal vein

which runs from the postnervulus to

the lower apex of the front wing of

wasps (fig. 31).

submarginal vein.—A longitudinal

vein paralleling the front margin at

the base of the front wing in some

wasps (fig. 57).

subspiracular protuberance.—A

swelling or projection below the

spiracular plates in puparia of some

species (fig. 65).

suture.—An external linelike groove

in the body wall.

tarsal claws.—A pair of small hooks

at the apex of the last segment on

the legs (fig. 40).

tarsus, pi. tarsi.—^The apical

segments of the leg, usually five in

number but sometimes less in

smaller wasps (fig. 34).

tergite.—^The dorsal (upper)

hardened plate of each abdominal

segment of wasps.

thorax.—^The middle of the three

main body segments to which are

attached the wings and legs. It is

comprised of the prothorax,

mesothorax and metathorax.

tibia.—^The fourth leg segment.

Typically the posterior of the two

longest leg segments, having one or

two spurs at the apex and narrower

than the femur (fig. 34).

38
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105.

44
Figures

Figures 1-29.—Tachinidae.

1. General wing of a tachínid.

2. General color pattern on thorax

of a tachinid (dorsal view).

3. General color pattern on thorax

of genus Nemorilla (dorsal view).

4. Diagrammatic dorsal view of a

tachinid.

5. Diagrammatic lateral view of a

tachinid.

6-9. Side view of heads ofCí/rcW/í/

(6), Eusisyropa (7), Nemorilla (8),

and Palexorista (9).

10-12. Semidiagrammatic figures of

compound eyes with hairs.

13-14. Semidiagrammatic figures of

side view of thorax to show strong

and weak pre-alar bristles (PRAL).

15-16. Semidiagrammatic figures of

humeral bristles.

17-20. Semidiagrammatic figures of

sternopleural bristles.

21-22. Semidiagrammatic figures of

marginal scutellar bristles.

23-25. Semidiagrammatic figures of

underside of abdominal segment 4

ofBlepharipa (23), Palexorista

inconspicua (24), and P. disparts

(25).

26. Semidiagrammatic figure of side

view of abdomen of females of

Compsilura and Blondelia.

27. Semidiagrammatic figure of

base of hind legs of typical Carcelia.

28-29. Semidiagrammatic figure of

median forceps (fused cerci) of

males ofExorista japónica (28) and

E. larvarum (29).

ACR = Acrostical bristles

AP = Apical cell

ANT = Antenna

AR - Arista

AS = Apicoscutellar bristle

B = Barette

DC = Dorsocentral bristles

DI = Discal cell

DS = Discoscutellar bristle

FR = Frontal bristles

FS = Frontal stripe

H = Humérus*

HA = Halter

HB = Humeral bristles

HPL = Hypopleuron*

lAL = Intra-alar bristles

IV = Inner vertical bristle

LS = Lateral scutellar

bristles

M = Media vein

m = Medial crossvein

MD = Median discal bristle

MM = Median marginal

bristle

MSNT = Mesonotum

MSNTS = Mesonotal suture

MSPL = Mesopleuron*

MSS = Mesothoracic

spiracle

MTPL = Metapleuron

MTS = Metathoracic spiracle

NP — Notopleuron*

NPB = Notopleural bristles

OC = Ocellar bristle

OV = Outer vertical bristle

PAC = Postalar callus

PAL = Postalar bristles

PFA = Parafacial area

PFR = Parafrontal area

PO = Proclinate orbital

bristles

POC = Postocular cilia

PPL = Propleuron*

PRAL = Prealar bristle

PSC = Postscutellum

PTPL = Pteropleuron*

PV = Postvertical bristle

R = Radius vein

45
r-m = Radio-medial

erossvein

RO = Reclinate orbital

bristles

SAL = Supra-alar bristles

SC = Scutellum

STB = Stigmatic bristles

STPL = Stemopleuron*

V = Vibrissa

W = Wing

WB = Wing base

* Bristle names correspond to

segment names

Figure 1.-

Figure 4.-

R4+5

M 1+2

Figure 2.—General Tachinid

Figure 3.—Nemorilla

46
Figure 5.—

Ro íy OV

Figure 6.— Figure 7,— Figure 8.-

Figure 9.— Figure 10.— Figure 11.— Figure 12.—

47
Figure 13.-

PRAL

Figure 14.

PRAL

Figure 15.— Figure 16.-

Figure 17.— Figure 18.- Figure 19.- Figure 20.—

Figure 21.— Figure 22. Figure 23

Figure 25. Figure 26 Figure 27.—

Figure 29.—

48
Figures 30-42.—Ichneumonidae

30. Mesochoms sp., front wing.

31. Phobocampe disparis (Viereck),

front wing.

32. Hyposoter tricoloripes

(Viereck), propodeum and first two

abdominal segments (dorsal view).

33. Coccygomimus disparis

(Viereck), propodeum and first two

abdominal segments (dorsal view).

34. Theronia atalantae fulvescens

(Cresson) (lateral view).

35. Casinaria sp., first abdominal

segment (lateral view).

36. Casinaria sp., head (facial

view).

37. Hyposoter tricoloripes

(Viereck), first abdominal segment

(lateral view).

38. Hyposoter tricoloripes

(Viereck), front wing.

39. Phobocampe disparis (Viereck),

first abdominal segment (lateral

view).

40. Theronia atalantae fulvescens

(Cresson), last tarsal segment.

41. Itoplectis conquisitor (Say),

head (facial view).

42. Coccygomimus pedalis

(Cresson), head (facial view).

AB—Abdomen

ABl—First abdominal segment

AB2—Second abdominal segment

ARE—Areolet

AS—Antennal sockets

ATI—First antennal segment

CB—Claw bristle

CL—Claw

CX3—Hind coxa

D—Discoideas

E—Eye

F3—Hind femora

FA—Face

GL—Glymma

HD—Head

LPN—Lower abscissa of

postnervulus

M—Médius

MD—Mandibles

MS—Mesopleural suture

N—Nervulus

O—OceUus

P—Petiole

PR—Propodeum

RVl—First recurrent vein

RV2—Second recurrent vein

SD—Subdiscoideus

SP—Spiracle

ST—Sternite

TA—Tarsus

TE—Tergite

TH—^Thorax

UPN—Upper abscissa of

postnervulus

Figure 30.

ARE

Figure 31.—

ARE

LPN

49
Figure 32.— Figure 33.

SP >|

V PR

> ABI

AB2

>PR

y ABI

> AB2

Figure 34.-

50
Figure 35.— Figure 37.-

MD

Figure 38.— Figure 39.—

Figure 40

51
Figures 43-50.—Braconidae

43. Apúnteles melanoscelus

(Ratzeburg), front wing

44. Meteorus pulchricornis

(Wesmael), front wing

45. M. pulchricornis (Wesmael),

propodeum and first abdominal

segment (dorsal view)

46. Rogas indiscretas Reardon,

propodeum and first abdominal

segment (dorsal view).

Al, R. indiscretas Reardon, head

(facial view).

48. Apanteles porthetriae

Muesebeck, anterior of abdomen

(dorsal view).

49. A. melanoscelas (Ratzeburg),

abdomen (dorsal view).

50. A. liparidis (Bouché), anterior

of abdomen (dorsal view).

AB I—First abdominal tergite

AB2—Second abdominal tergite

AB3—^Third abdominal tergite

ATI—First antennal segment

Cl—First cubital cell

C2—Second cubital cell

C3—Third cubital ceU

CL—Clypeus

CU—Cubitus

E—Eye

FA—Face

ICI—First intercubitus

IC2—Second intercubitus

MD—Mandible

PR—Propodeum

R—Radius

RVl—First recurrent vein

ST—Stigma

Figure 43.— Figure 44.-

52
Figure 45. Figure 46.

> PR

> ABI

> ABI

Figure 47.- Figure 48.-

MD

Figure 49.- Figure 50.

ABI

AB2

53
Figures 51-62.—Chalcidídae,

Encyrtidae, Eupelmidae, Scelíonídae,

and Torymidae.

51. Brachymyeria sp., head (lateral

view).

52. Brachymeria sp., hind leg.

53 Anastatus disparis Ruschka,

front wing.

54. Telenomus sp., front wing.

55. Monodontomems sp. (lateral

view without legs).

56. Ooencyrtus kuvanae (Howard)

(lateral view without antennae and

legs).

57. O. kuvanae (Howard), front

wing.

58. O. kuvanae (Howard), thorax

(dorsal view).

59. Anastatus disparis Ruschka,

male, thorax (dorsal view).

60. A. disparis Ruschka, female,

thorax (dorsal view).

61. Scelionidae, thorax without

appendages (lateral view).

62. Chalcidoidea, thorax (lateral

view) (diagrammatic).

AX—Axilla

CT—Coxal tooth

CXI—Front coxa

CX2—Middle coxa

CX3—Hind coxa

E—Eye

F3—Hind femora

EC—Frontal carina

EG—Femoral groove

GM—Genotemporal margin

MN—Mesonotum

MP—Mesopleuron

MV—Marginal vein

NE—Notauli

PC—Postorbital carina

PR—Pronotum

SC—Scutellum

SM—Submarginal vein

SV—Stigmal vein

T3—Hind tibia

TE—^Tegula

Figure 51.

Figure 52.

CT CX3

Figure 53.—

MV SV

Figure 54.—

MV SV

54
Figure 55.— Figure 56.-

CXl MP CX3

Figure 57.-

^CX2

CXI CX3

Figure 58.-

MN

AX

Figure 59.—

MN

AX

Figure 60.-

MN

AX

Figure 61.

55
Figures 63-72.—Puparia of

Tachinidae (on left, posterior end; on

right, spiracular plates, subspiracular

protuberance, and distance from anus).

63. Parasetigena silvestris

64. Palexorista inconspicua

65. Blepharipa pratensis

66. Compsilura concinnata

67. Careelia amplexa

68. Lespesia aleîiae

69. Lespesiafrenchii

70. Eusisympa viritis

71. Nemorilla pyste

72. Blondelia nigripes

SP—Spiracular plate

SS—Spiracular slit

SSP—Subspiracular protuberance

A—Anus

Figure 63.-

^i^U

Figure 65.

Figure 66. Figure 67.

0<

SSP SSP

o

56
Figure 68.— Figure 69.—

Figure 70.— Figure 71.—

O.

Figure 72.-

57
Figures 73-75.—Cocoons of

Ichneumonídae

73. Meîeoms pulchricorn is

Wesmael

74. Hyposoter îricolonpes (Viereck)

75. Phobocampe disparis (Viereck)

Figure 73.—

Figure 74.—

Figure 75.—

58
Acknowledgments

We are indebted to many colleagues

for assistance and information in the

preparation of this handbook. Our

special appreciation goes to Robert

Carlson, Eric Grissell, Paul Marsh,

and Carl Muesebeck for their many

helpful suggestions and courtesies

during visits to the Systematic

Entomology Laboratory, U.S.

Department of Agriculture, at the

U.S. National Museum,

Washington, D.C. We thank the

Entomological Society of America

for permission to use several parts

of the text and figures of "Tachinid

parasites of the gypsy moth,

Lymantria dispar, with keys to

adults and puparia,'' by Sabrosky

and Reardon, 1976, Miscellaneous

Publication 10(2). Most of the

drawings of the wasps were made

by Joan Clark, Pennsylvania

Department of Environmental

Resources; drawings of the

tachinids were done by Kathleen

Schmidt, University of Connecticut.

We also thank R.W. Carison, J.R.

Coulson, P.B. Dowden, E. Grissell,

RM. Marsh, C.W. Sabrosky and K.

Welch for review of this manuscript.

i^ U.S. GOVERNMENT PRINTING OFFICE: 1979—273-083

59