STUDIES ON THE HYMENOPTERA

Henry K. Townes has been an active student of the Hymen-

optera for the past nearly fifty years. His main love has been

the study of the Ichneumonidae, but he has worked on a

number of other families of the Hymenoptera and Diptera. His

contributions in the Ichneumonidae in the form of taxonomic

revisions of the North American fauna, Catalogs and reclassi-

fications of the Ichneumonidae of various zoogeographic regions,

and the monographs on the Genera of the Ichneumonidae of the

World, have given us fundamental literature which is guiding

the research activities of the beginner as well as the advanced

students of the Hymenoptera throughout the world.

The articles presented in this commemorative volume have

been specially written by his students and colleagues to honor him

for his achievements on the occasion of his seventieth birthday on

January 20, 1983. This has been done in a very short time and

certain inconsistencies in the style of presentation, etc. could not

be rectified.

We wish Dr. Henry K. Townes a long and productive life and

hope to see many more of his contributions on the Hymenoptera.

STUDIES ON THE HYMENOPTERA

TABLE OF CONTENTS

Henry K. Townes -- The Man and the Scientist

Scientific publications of Henry K. Townes

Description of the Townes Collection of Hymenoptera

Dedication

10.

12.

13.

15.

Denis F. Owen: A hole in a tent or how to explore insect

abundance and diversity .

Mattias Idar: A new Nearctic species of Hadrodactylus.

W. R. M. Mason: A new South African subfamily related to

Cardiochilinae (Hymenoptera: Braconidae).

Lars Huggert and Lubomir Masner: A review of Myrmecophilic

symphilic Diapriid wasps in the Holarctic realm, with descrip-

tions of new taxa and a key to genera (Hymenoptera: Procto-

trupoidea: Diapriidae).

C. M. Yoshimoto: A new genus of Tetrastichinae from New

world (Hymenoptera: Chalcidoidea: Eulophidae) .

M. J. Sharkey: Marjoriella, a new Neotropical genus of

Agathidinae (Hymenoptera: Braconidae) .

Klaus Horstmann: Revision of species of Western Palearctic

Ichneumonidae described by French authors.

D. R. Kasparyan: A new Eastern Palearctic genus of the

subfamily Phrudinae (Hymenoptera: Ichneumonidae) .

M. Capek: Relationships and taxonomic placement of the

genus Blacus (Hymenoptera: Braconidae) .

B. R. Subba Rao and M. Hayat: Key to the genera of

Oriental Mymaridae, with a preliminary catalog

(Hymenoptera: Chalcidoidea).

Rolf Hinz: The biology of the European species of the genus

Ichneumon and related species (Hymenoptera: Ichneumonidae).

G. van Rossem: A revision of Western Palearctic Oxytorine

genera. Part Ill. Genus Proclitus (Hymenoptera: Ichneumon-

idae).

P. M. Sanborne: Two new Palearctic species of Sesioplex

Viereck (Hymenoptera: Ichneumonidae) including phylogeny,

zoogeography and a key to the world species.

Virendra Gupta: A review of the world species of Dyspetes

(Hymenoptera: Ichneumonidae) |

T. Tachikawa: A new species of Astichus parasitic on a

cid beetle from Borneo (Hymenoptera: Eutophidae)

101

116

119

125

+51

153

166

Tf7

189

150

152

165

176

188

190

16.

re.

18.

20.

at.

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23.

24.

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27.

28.

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30.

St.

32.

33.

34.

Contrib. Amer. Ent. Inst., vol. 20, 1983

|. D. Gauld and G. A. Holloway: A new genus of endaseine

Ichneumonidae from Australia (Hymenoptera). 191

L. Mochar: Two interesting Sulcomesitus species from

Africa (Hymenoptera: Bethylidae). 198

Reijo Jussila: Addition to the revision of the genus Atractodes

(Hymenoptera: Ichneumonidae) of the Western Palearctic Region.201

Charles C. Porter: A new cloud forest Trachysphyrus from

Ecuador and Colombia (Hymenoptera: Ichneumonidae). 205

Karl-Johan Hedqvist: A new species of the genus Karpinskiella

(Hymenoptera: Chalcidoidea: Pteromalidae). 213

Franco Frilli: Studies on Hymenoptera: Ichneumonidae. |X.

The Rudolf Jussel's Collection. 218

Maximilian Fischer: Descriptions of some new African species

of Ophius from the Townes collection (Hymenoptera:

Braconidae: Opiinae). 221

J. Papp: Three new Opiinae species from India (Hymenoptera:

Braconidae). 2c?

Santosh Gupta: A revision of the genus Prosthoporus

(Hymenoptera: Ichneumonidae). 245

Michael G. Fitton: A new species of Bioblspsis (Ichneumon-

idae: Diplazontinae) from North America. 254

Alexender P. Rasnitsyn: Ichneumonoidea (Hymenoptera) from |

the Lower Cretaceous of Mongolia. 259

Josef Sedivy: Tersilochinae as parasitoids of insect pests of

winter rape (Hymenoptera: Ichneumonidae). 266

Robert A. Wharton: Senwot, a new genus of Alysiinae

(Hymenoptera: Braconidae) from Africa. 217

C. van Achterberg: A revision of the new tribe Brulleiini

(Hymenoptera: Braconidae). 281

Mark R. Shaw: On evolution of endoparasitism: The biology

of some genera of Rogadinae (Braconidae). _ 307

Jean Paul Aeschlimann: Notes on the variability of Microctonus

aethiopoides Loan (Hymenoptera: Braconidae: Euphorinae). 329

David Rosen and Amos Alon: Taxonomic and biological studies

on Diversinervus cervantesi(Girault) (Hymenoptera: Encyrtidae),

a primary parasite of soft scale insects. 336

Paul Marsh. A taxonomic study of the South American Megalo-

proctus Schulz (Hymenoptera: Braconidae: Doryctinae). 363

E. Diller: Eine neue art der gattung Herpestomus Wesmael

aus Asien (Hymenoptera: Ichneumonidae: Phaeogenini). 381

380

383

35.

36.

oT.

38.

og.

40.

E. Haeselbarth and Conrad Loan: Townesilitus, a new genus for

a species group in Microctonus (Hymenoptera: Braconidae,

Euphorinae) .

Conrad Loan: Host and generic relations of the Euphorini

(Hymenoptera: Braconidae).

Vinalto Graf: Ichneumofauna do Sudeste e sul do Brasil. V.

Nova especie de Philodrymus (Anomaloninae, Hymenoptera).

Luis De Santis: Insectos que destruyen el expartillo (Stipa

brachychaeta) y sus parasitoides.

Robert W. Matthews and Janice R. Matthews. Malaise trap:

The Townes model catches more insects.

Biological studies on Goniozus legneri Gordh (Hymenoptera:

384

388

398

402

428

Bethylidae), a primary external parasite of the naval orangeworm

Amyelois transitella and pink bollworm Pectinophora gossy-

piella (Lepidoptera: Pyralidae, Gelechiidae).

New taxa described in the volume

433

469

387

397

401

427

432

468

471

HENRY K. TOWNES, Jr.

The Man and the Scientist

An Appreciation

Virendra Gupta

My association with Henry Townes goes back to 1957, when | came to

work as a graduate student under his guidance. Ever since | have admired his

personal and professional qualities and have developed an appreciation for him

as a man and a scientist, unique in many ways.

When | began my work on the Ichneumonidae of India in 1954, | soon

discovered how inadequate the existing literature was. During the course of

my correspondence with the ichneumonid workers, | came in touch with Henry

Townes, who helped me with literature, specimens, and advice. This culmi-

nated in my coming to the University of Michigan as a graduate student in

[957 to work on the Ichneumonidae. That was a turning point in my career

and the beginning of an association with a leading taxonomist, which | have

always cherished. Before leaving India for Ann Arbor, | was advised that the

American Professors rarely see their graduate students, at best once a week,

and that | should not bother him with problems (as | used to do in India when

| did not get anywhere). To my surprise, Henry was often by the side of my

desk, looking at my specimens and giving me practical tips. | soon discovered

that he was freely accessible and that | need not wait for the weekend to

solve my queries. When | returned to work with him in 1980, after a gap of

nearly 19 years, | found his working habits the same. He was by the side of

my desk as often as he got tired of looking into his own microscope. During

the intervening years | was in active correspondence with him and he readily

offered me help and advice on all taxonomic matters.

| was no exception. Henry has been most communicative and helpful to

all his students and colleagues. His attitudes have gone a long way towards

progress in ichneumonid taxonomy that we see all over the world. He at-

tracts students from all over the world who come to study his collections and

seek his advice.

THE FORMATIVE YEARS

Henry Keith Townes, Jr. was born in Greenville, South Carolina on |

January 20, 1913, the third of the six children of Henry K. and Ellen Hard

Townes. According to his family, he has been sensitive, independent, self-

contained, mischievous, and a naturalist from his very early age. His family

encouraged his interests. He was unusually knowledgeable about natural

sciences for his age, much to the annoyance of some of his teachers.

Henry Townes' first collecting activity was the butterflies. He set out

to collect all the species of butterflies of South Carolina. »At the age of 10

or Il he won a second prize for his exhibit of 47 out of 50 kinds of butter-

flies that were known to occur in South Carolina. He missed the first prize,

not for the species that he had not collected, but because his contender had

arranged and prepared the butterflies better. From the butterflies his

2 Contrib. Amer. Ent: inst... vol. 20, 1983

interest shifted to the crane-flies, when he came in contact with leading

taxonomists like J. Speed Rogers and C. P. Alexander.

Henry Townes took his college training at Furman College (now Furman

University) in Greenville, South Carolina, from 1929 to 1933. He completed

the requirements for a B.S. degree in three years. His family felt that he

was too young to leave for a graduate school; so during his fourth year at

Furman he took courses in Languages for a B.A. degree. At the 1933 com-

mencement excercises he was the only graduate to receive two degrees --

B.S. and B.A. When his brother Charles, repeated this achievement, the col-

lege made a rule that a student could not be awarded more than one degree

at a commencement.

GRADUATE WORK

Henry Townes joined the Cornell University in the fall of 19383 to study

Entomology. Cornell was perhaps the best place to study Entomology at that

time. The real challenge and excitement must have come from his courses

in Entomology with well-known and Knowledgeable professors. After some in-

vestigation of smaller Hymenoptera he decided to do his thesis on some tax-

onomic problem in the Ichneumonidae under Professor J. Chester Bradley. This -

was a challenging position, for Professor Bradley was known to be an exacting

scholar and to leave his students pretty much on their own. Before Henry

only J. D. Hood had completed his doctorate under Professor Bradley.

In 1933 there was very little reliable literature on the Ichneumonidae,

nothing to provide a really practical introduction to the family. There was

the publication of W. C. Davis on the subfamily Tryphoninae. With this as

a guide, Henry tried to classify the Cornell collections. He found the going

difficult and, while some of the specimens before him could be identified, many

could not. There seemed to be gaps, and confusions and he was left with many

questions. Eventually it became evident to him that the only way to provide

reliable answers to the questions was to check the type-specimens. So Henry

began to travel to see the type-specimens and comparing his collections with

the types and labeling them as homotypes. It was soon clear that there were

many misdeterminations in the literature and many synonymies and the best

way to go about was to revise groups based on a study of the types. Prior

to him not much attention had been paid to the study of the types. The

literature was supposed to be sacrosanct and provide all the information neces-

sary for identification. What did not match with the literature was supposed-

ly a new find. Henry initiated the study of the types -- which is now a

standard practice in any taxonomic study.

Henry continued to work towards his Ph.D. degree, supporting himself

with odd jobs, building up his own collections, and comparing them with the

types. In 1937 he finished his thesis on 'The Nearctic species of Netelia",

in which many new innovations were made about the classification of the

Ichneumonidae. During the summers of his graduate work he was employed

by the New York State Stream Survey, which also gave him an opportunity

to make collections of his own. This work and the aquatic collecting also

resulted in his publication on the Nearctic Tendipedini.

Henry K. Townes 3

MARJORIE CHAPMAN

Henry Townes and Marjorie Chapman, a native of Westerly, Rhode

Island, a graduate of Mt. Holyoke College and with a Ph.D. in Botany from

Cornell in 1935, were married on October 9, 1937.

They had met at Ithaca at the home of the Baptist student pastor where

she was preparing dinners and baby sitting to help pay her expenses. Their

courtship included collecting trips, visits back and forth to Mt. Holyoke

College where she taught for two years and to meet the two concerned fam-

ilies. Henry very much admired the close cooperation between the Comstocks

(John Henry and Anna B.) and Marjorie agreed that a similar partnership

would be an ideal to strive for. They have worked together ever since. Her

contributions towards the advancement of Ichneumonology, though cryptic, are

as important as the contributions of Henry. She sacrificed her career in

Botany for the cause of ichneumonid taxonomy and has willingly and readily

taken care of all the chores that go with research. They have inspired me

in the same way as Comstocks inspired them.

THE CAREER

In spite of his academic accomplishments, Henry had a hard time finding

a job, particularly at the time when he was planning to get married. Those

were the days of the recession. After considering almost any possibility, even

working on the Cornell grounds crew, he was appointed to a first semester

position in the Zoology Department at Syracuse University with a salary of

$800 for the semester. This was later extended through the full school year.

In 1938, they returned to Ithaca where Henry had been appointed to teach

medical entomology to substitute for Professor Robert Matheson who was on

leave. Adding to his collections was a matter of primary importance to Henry

and in Ithaca and elsewhere, both collected ichneumons and other insects when-

ever time and opportunities were available. As Henry became known for his

ichneumonid works, gifts of specimens collected by friends and collections for

naming started coming. These supplemented their personal collectings and the

collections continued to grow. At the same time they maintained and worked

upon a card file on the Ichneumonidae, which was continuously updated as and

when the type-specimens were examined and synonymies and homonymies dis-

covered. With the return of Professor Matheson, the position at Cornell ter-

minated. The next opportunity came with a grant to work at the Academy

of Natural Sciences of Philadelphia preparing "A catalog and reclassification

of the Nearctic Ichneumonidae". This was completed in 1941, and after usual

publication dalays, published in 1945-46.

The preparation of the Nearctic Catalog required hundreds of decisions

about nomenclature. It was obvious that such an abundance of decisions must

be handled objectively rather than subjectively. It was necessary for Henry

to review his options and define his rules. It became clear that the only way

to reach the desired nomenclatural stability was to apply the rule of priority

without exception, without presenting cases to the International Commission

for rulings and that could be overturned with a different set of commissioners

persuaded by taxonomists who had different reasons for preserving other names.

Henry has ardently followed the Law of Priority, much to the annoyance of

many workers. However, this practice has produced a nomenclature in the

4 Contrib. Amer. Ent. Inst., vol. 20, 1983

Ichneumonidae that has come to stay and has already gained acceptance with

the majority of workers.

Henry has always given the highest priority to collecting of specimens

afresh for any taxonomic revision. Both Henry and Marjorie have been active

collectors and have undertaken several collecting trips. Early in their career

they collected several thousand specimens in the Pacific northwest and in

Western U.S.A. These collections yielded valuable specimens from areas that

have not been previously explored thoroughly. The taxonomic revisions pro-

duced at a later date incorporating all these collections have given us a better

picture of the nearctic fauna and a better understanding of the faunal rela-

tionships.

With the expiration of the catalog grant, there was again the need to

find a paying job. The family, now with a son and a daughter, was left in

South Carolina while Henry worked briefly in Virginia on insects itn apple or-

chards. Later he moved to the U.S. National Museum, as a taxonomist in

Orthoptera. They settled in Takoma Park, a suburb of Washington, which is

documented by the fact that there are thousands of ichneumonids from Takoma

Park in his collections. The Pearl Harbor had just been attacked. As more

and more taxonomists were drafted for the war service, work on the wasps

and on the Ichneumonidae were added to Henry's job. It was during this time

that he wrote papers on Dermaptera and on the Pepsinae and Ceropalinae in

the Psammocharidae (= Pomplidae). He also initiated the project to write a

catalog of Nearctic Hymenoptera on the pattern of the Ichneumonid catalog.

With several of his colleagues at the National Museum and with the collabora-

tion of a number of other specialists, this catalog was completed and published

in 1951. It was during this time that DDT was beginning to be used on a broad

scale to control insect pests, that Henry spent a summer on a team surveying

the effects of spraying DDT on the insects in a forest in Central Pennsylvania,

one of the earliest efforts to evaluate the consequences of such a treatment.

When R. A. Cushman retired, Henry was formally appointed to the National

Museum position as a specialist on the Ichneumonidae.

Henry was apparently never satisfied with his job at Washington. His

views on taxonomy and nomenclature conflicted with those of his colleagues.

He also did not like to live far away from his working place. In 1949 the

Townes moved to Raleigh, North Carolina. Henry Townes accepted a position,

at a lower salary, at the North Carolina State University to work on insects

affecting tobacco. His work on the Ichneumonidae slowed down, but his col-

lecting continued, with all his spare time and weekends devoted to that. There

were accomplishments in Economic Entomology: Perhaps the most notable was

the initiation of studies on the role of Polistes wasps in biological control

in tobacco fields which led to various studies on the biology and behavior of

those insects. His contacts with the students were stimulating, both for him

and his students.

EXQTIC COLLECTIONS

Henry's first experience of collecting exotic fauna came in 1945 with

his appointment as an Entomologist on a survey of the Mandated Islands of

the Pacific. The specimens from this survey were deposited in the National

Museum, but the experience on the trip was a step in expanding his view on

the world fauna. The next opportunity came with his appointment as an

advisor to the Philippine Government on pests of rice and corn with the

Henry K. Townes 5

Economic Cooperation Authority. From 1952 to 1954 the Townes family was

resident in Manila. While attending to his job, he utilized every possible

opportunity to collect insects in the Philippines particularly the Hymenoptera.

A very valuable lot of exotic specimens was added to the Townes Collection.

This was augmented on the way back to the U.S.A. in 1954 with a stop in

Japan to study the types in Sapporo and to collect in the environs of that city

and in the Japanese alps.

His collecting in the Philippines had a direct effect on his interest in

the Oriental fauna. This enabled several students from the Orient to work

with him, notably Clare Baltazar, Sui-Chen Chiu, Virendra Gupta and Setsuya

Momoi. The Townes have collected the Ichneumonidae practically all over the

world when they traveled to study types for their catalogs and revisions of

the genera of the Ichneumonidae. Where ever they went, they left a lasting im-

pression with the students of Hymenoptera. Interest in the taxonomy of the

Ichneumonidae thus flourished throughout the world.

A BREAK

Henry Townes has always been restless and looking for opportunities to

work on the Ichneumonidae and other parasitic Hymenoptera. The opportunity

finally came in 1956 with the Dow Chemical Company awarding him a grant

to work on the Ichneumonidae at the University of Michigan for a period of

five years. He resigned his permanent job at Raleigh and moved to Ann Arbor,

Michigan. This was made possible by the late R. R. Dreisbach, a friend and

collaborator with the Townes, who was a chemist at the Dow Chemical Com-

pany. Dreisbach, who was himself a student of the Aculeate Hymenoptera,

conceived the idea and used his influence to get the grant, with no prospect

of any profitable gain for the Dow Chemical Company. That grant was there-

fore unique for a chemical company to support research unconnected with

their activities. It also provided the break that the Townes were looking for

to devote their undivided attention to the study of the Ichneumonidae.

During this period the launching of the Sputnik was a blessing in disguise

for the scientists in the country. Scientific research was revitalized with

various Governmental agencies providing funding for basic research. The

National Science Foundation provided support for ichneumonid taxonomy and

for cataloging the Ichneumonidae of various zoogeographical areas. The

National Institutes of Health also provided grants to work on the Ichneumoni-

dae and on the Genera of Ichneumonidae of the World. There was freedom

for work and research on the taxonomy of the Ichneumonidae flourished.

THE AMERICAN ENTOMOLOGICAL INSTITUTE

As work on the Ichneumonidae progressed and collections grew, the

Townes faced two problems: a publication outlet for their voluminous works

in the style they liked, and adequate space and housing facilities for their

collections. These were solved by the Townes by establishing a non-profit

research institution, the "American Entomological Institute" in 1962, and build-

ing their own laboratory for research in 1964. They also started their own

publications, the "Memoirs"! and the "Contributions" of the American Entomol-

ogical Institute. To date 34 volumes of the Memoirs and 19 volumes of the

Contributions have been published, including works of various authors as well

as their own. Research has been conducted under various grants to the

6 Contrib. Amer. Ent. Inst., vol. 20, 1983

Institute from the NSF and NIH. The Townes have also worked without ex-

ternal financial support. Taxonomic work has continued uninterrupted, re-

gardiess of whether a grant was available or not. Several collecting trips have

been undertaken by them. The Townes Collection has grown, now totaling

over 793,000 pinned and labeled specimens. An excellent library of Hymenop-

tera has been built up. This self contained research center has all that a

working taxonomist would want. It attracts a number of scholars every year

from far and near. It is a rewarding experience to work there, for Henry's

help and advice are freely available. World collections are there to examine,

and above all, the hospitality of the Townes is unforgettable. Marjorie

Townes especially takes good care of her guests, providing them with excel-

lent meals. One could walk in any time and will be well cared for!

Significant advancements have been made by the Townes and their stu-

dents in the study of the Ichneumonidae during the past 25 years or so. The

Townes have published revisions of the Ichneumonidae of North America,

Catalogs and reclassification of Ichneumonidae of various zoogeographical re-

gions, monographs on the World genera of the Ichneumonidae, a monograph

on the Serphidae of the World, and miscellaneous papers on ichneumonid tax-

onomy and nomenclature. His students and colleagues have also been active,

either collaborating with him or working independently on ichneumonid fauna

of different parts of the world. Notable among them are Baltazar, Chiu,

Gupta, Momoi, Dasch, and colleagues like Carlson, Porter, Gauld, Fitton,

Aubert, Horstmann, Oehlke, Hinz, van Rossem, Zwart, and several others. We

thus have solid basic literature on the Ichneumonidae on a world-wide basis,

which does not appear to exist for any other group of insects. Further re-

search and revisions on a world wide basis is Wiearee a: And credit for this

goes to the Townes.

THe. FINALE

One of the things that had bothered the Townes for the past few years

is the future of the collections and the Institute. Having labored so hard to

build up the world's best collection of the Ichneumonidae and an excellent

collection of other groups of the Hymenoptera, one is naturally inclined to

find a satisfactory place where the collections will be well cared for. More

than that the Townes are very much concerned with the lack of support for

taxonomic research in this country. While most of the established institutes

would like to possess his collections, no one was ready to insure that tax-

onomic research will continue uninterrupted on Ichneumonidae and other para-

sitic Hymenoptera. His efforts in finding a suitable locale for his collections

and taxonomic research culminated in the establishment of the "Center for

Parasitic Hymenoptera" in 1982 at the University of Florida, Institute of Food

and Agricultural Sciences, thanks to the vision of Professor K. R. Tefertiller,

Vice-President for Agricultural Affairs at the University of Florida and the

Chief of the IFAS. This Center for taxonomic research on the parasitic

Hymenoptera is unique in the World. The Townes collections and library forms

the nucleus of the Center, and the Center will become the focal point of tax-

onomic research and training in the field of parasitic Hymenoptera. It is a

tribute to Henry Townes, and his vision, who has relentlessly worked towards

advancement of taxonomic research on the parasitic Hymenoptera.

The Townes 7

THE IMPACT

Scientific research needs dedication, whether it is taxonomy or molec-

ular biology. Some may be more fashionable than the others, but all are

equally demanding. In certain ways taxonomic research needs more dedica-

tion, because it takes a life time to specialize in one group of insects. No

one person can specialize in all insects. The Townes have dedicated them-

selves to the study of Ichneumonidae. Although they have their idiosyncra-

sies, as any other scientist would have, they have given us several directions

for research. Extensive collections have always been emphasized by them

before undertaking any taxonomic revision. The study of types has been shown

to be of paramount importance in all revisionary studies. Cataloging has been

shown to be original research rather than merely copying data from zoological

records. All the catalogs published by Townes and his students contain ori-

ginal information on the types and keys to genera. The taxa have been

taxonomically assessed after the study of the types, resulting in several new

combinations, synonymies, homonymies, etc. Without such catalogs further

research on taxonomic revisions was impossible, as many species were origi-

nally assigned to wrong genera. With no keys to genera existing before, it

was impossible to progress in the right direction. All his revisions included

Original illustrations at a time when illustrations were not fashionable, par-

ticularly in the U.S.A. The most useful tools that they have given us, are the

volumes on the Genera of the Ichneumonidae, with each genus illustrated,

keyed and described. What else does a beginner or an advanced student need

to continue work on Taxonomy?

The Townes are readily accessible. Their hospitality is something to

remember and working with them something to cherish. It is only befitting

that we, his students and colleagues, celebrate the 70th birthdate of: Henry

Townes by presenting this collection of articles on Hymenoptera to him as a

token of our regards and affection for him and his wife and working partner,

Marjorie Townes.

10.

13.

14.

1936.

[9o7.

1938.

fon8.

1939:

1939.

1940.

1943.

Contrib. Amer. Ent. Imst.; vol. 20, 1983

SCIENTIFIC PUBLICATIONS OF HENRY K. TOWNES, ‘JR.

(with F. R. Shaw). A preliminary report of the Mycetophilidae

of North and South Carolina. Bull. Brooklyn Ent. Soc. 3l: 204-208.

(with F. R. Nevin). A study of the larger invertebrate forage

organisms in selected areas of the Delaware and Susquehanna

watersheds. Supplement to the twenty-fifth annual report of the

New York State Conservation Department. 1936, pp. 195-204 (pp.

201-204 by Townes).

(with F. R. Nevin). Studies of invertebrate forage organisms in

selected areas with notes on the effects of pollution upon them.

Supplement to the twenty-sixth annual report of the New York

State Conservation Department. 1937, pp. 214-227.

Studies on the food organisms of fish. Supplement to the twenty-

seventh annual report of the New York State Conservation Depart-

ment. 1938, pp. 162-175.

Ichneumon hibernation in northeastern United States (Hymenoptera,

Ichneumonidae). Ent. News 49: 219-221.

Pammegischia and Trichofoenus discarded (aulacoid Hymenoptera).

Canad. Ent. 70: 254-255.

The Nearctic species of Netelia (Paniscus of authors) and a revi-

sion of the genera of Neteliini (Hymenoptera, Ichneumonidae).

Lloydia |: 168-231.

Protective odors among the Ichneumonidae (Hymenoptera). Bull.

Brooklyn Ent. Soc. 34: 29-30.

The correct generic position and synonymy of Symphobus pleuralis

(Cresson) and of Plectiscus pleuralis Provancher (Hymenoptera).

Canad. Entice Oe.

Corrections to the Gahan and Rohwer lectotypes of Provancher's

Ichneumonidae (Hymenoptera). Canad. Ent. 71: 91-95.

Ecological studies on the Long Island marine invertebrates of

importance as fish food or as bait. Supplement to the twenty-

eighth annual report of the New York State Conservation Depart-

ment. 1939, part | of the salt water section, pp. 163-176.

Notas sobre Ichneumonidae Venezolas. Bol. Soc. Venezol. Cienc.

Nat. 5: 299-30,

A revision of the Pimplini of eastern North America (Hymenop-

tera, Ichneumonidae). Ann. Ent. Soc. Amer. 33: 283-323.

A key to the families of Hymenoptera and a general account of

sixteen families of parasitic Hymenoptera. In Comstock, J. H.:

An Introduction to Entomology. Ninth edition revised, 1940 (pp.

891-894, 917-9384, 939-949). Pub. by the Comstock Pub. Co.,

Ithaca, N.Y.

(Book review). Outlines of Entomology by A. D. Imms. Proc.

Ent. Soc. Wash. 45: 95-96.

16.

18.

19.

20.

ei,

3 ae

23.

24.

20+

20.

29.

30.

31.

oe.

33.

1944.

1944

1945.

1946.

1947.

1948.

Henry K. Townes 9

Some taxonomic and biological observations on the chironomids.

Proc. Ent. Soc. Wash. 46: 163-164 (in minutes).

and 1945. A catalogue and reclassification of the Nearctic

Ichneumonidae (Hymenoptera). Mem. Amer. Ent. Soc., No. Il, in

2 parts, 925 pp.

The Nearctic species of Tendipedini [Diptera, Tendipedidae

(= Chironomidae)]. Amer. Midland Nat. 34: I-206.

(Book review). The Aphidiinae of North America (Braconidae:

Hymenoptera), by Clyde F. Smith. Amer. Midland Nat. 33: 796-

797.

(with M. C. Townes). A key to the genera of Eclytini

(= Thymaridini) and a revision of the Neotropical species (Hymen-

optera, Ichneumonidae). Bol. Ent. Venezolana 4: 41-53.

A list of the genera and subgeneric names of Dermaptera with

their genotypes. Ann. Ent. Soc. Amer. 38: 343-356.

Tendipedidae, in Stuardo: Catalogo de los dipteros de Chile, pp.

42-47. Ministerio de Agricultura, Santiago.

Entomological results from the treatment of large areas with DDT.

Minutes of 4th meeting of subcommittee on Dispersal, National

Research Council Insect Control Committee, pp. 31-34.

(with C. H. Hoffman, R. |. Sailer, and H. H. Swift). Field

studies on the effect of DDT on aquatic insects. "E" series of

U.S. Dept. Agriculture, No. 702, 28 pp.

Entomological definitions, in Funk and Wagnall: New Practical

Standard Dictionary.

Two earwigs new to the United States. Proc. Ent. Soc. Wash. 48:

47 (in minutes).

The generic position of the Neotropic Ichneumonidae (Hymenop-

tera) with types in the Philadelphia and Quebec museums,

described by Cresson, Hooker, Norton, Provancher, and Viereck.

Bol. Ent. Venezolana 5: 29-63.

(Book review). Insect Dietary, by C. T. Brues. Proc. Ent. Soc.

Wash. 48: 131-132.

A Eumenes wasp and six adventive Ichneumonidae new to Hawaii

(Hymenoptera). Proc. Hawaiian Ent. Soc. 13: 105-106.

Results of entomological inspection tour of Micronesia. Published

by the Economic Survey of the U.S. Commercial Company,

Honolulu. 53 pp.

The serphoid Hymenoptera of the family Roproniidae. Proc. U-S.

Natl. Mus. 98: 85-89.

Czechoslovakian insect pins are now available. Ent. News 59: 75-

76.

Pyrex bottles for potassium or sodium hydorxide solutions. Proc.

Ent. Soo. Wash. 50:85.

34.

35.

36.

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38.

39.

AQ.

Al.

42.

43.

AA.

45.

46.

47.

48.

49.

50;

1948.

1949.

1950.

1951.

18S!.

1952.

LO52.

1952.

[952

1952,

952.

Contrib: Amer. Gat. inst. vol, 20, 1983

Field work on the Ichneumonidae (Hymenoptera). Amer. Philos.

Soc. Yearbook 1947: 148-149.

(with: C. He Hoffman, H. H. Swift, and Pel. Sailer). Field

studies on the effects of airplane applications of DDT on forest

invertebrates. Ecol. Monogr. I9: I-46.

Collecting Ichneumonidae (Hymenoptera) in California and

Colorado. Amer. Philos. Soc. Yearbook. 1948: 132-134.

The Nearctic species of the family Stephanidae (Hymenoptera).

Proc. Us. Natl. Mus. 99: 3G6)-370.

The Nearctic species of Evaniidae (Hymenoptera). Proc. U.S.

Natl. Mus. 99: 525-539.

The effectiveness of DDT against dermestids in insect boxes.

Proc. Ent... Soc. Wash... 5!l? 165-168.

A pest that's a pushover (nemocerous larvae in tobacco plant

beds). Research and Farming. Autumn, 1950, p. 6.

(with M. Townes). A revision of the genera and of the American

species of Tryphonini (Hymenoptera: Ichneumonidae). Ann. Ent.

Soc. Amer. 42: 321-447.

The Nearctic species of Gasteruptiidae (Hymenoptera). Proc. U.S.

Natl. Mus. 100: 85-145.

Tobacco insect contro! in North Carolina. (N.C, Agric. Expt. Sta.

Extension Circ. no. 351, 6 pp. (also, a revised edition, 1951).

A cabinet for Schmidt boxes. Coleopterists' Bull. 5: 21-27.

(with M. Townes, C.F.W. Muesebeck, K. V. Krombein, and

others). Hymenoptera of America North of Mexico-Synoptic

catalog. U.S. Dept. Agr. Agr. Monog. no. 2, 1420 pp. (pp. 89-90,

184-410, 655-661, 907-909, 9II-921, and 939-940 by Townes).

(with M. Townes). A revision of the genera and of the Nearctic

species of Grypocentrini (Hymenoptera, Ichneumonidae). Proc.

Ent. Soc. Washington 53: 30l-3i3.

Tribe Tendipedini, in The Diptera or true flies of Connecticut:

Fifth fascile: Midges and gnats. Bull. Conn. Geol. Nat. Hist.

Surv. 80: 27-147.

[Nomenclatural corrections on Puerto Rican Ichneumonidae]. Jour.

Agr. Univ. Puerto Rico 32: 765.

TDE insecticide emulsions for tobacco. N.C. State College Ent.

Leaflet no. 5l, | p.

Benzene hexachloride (BHC) and lindane on tobacco. N.C. State

College Ent. Leaflet no. 56, 2 pp.

(with Henry Howden). On the type species of the genus

"Geotrupes" Latreille, 1796 (class Insecta, order Coleoptera) and

a discussion of the Fabrician (1898) usage of other names proposed

by Latreille in 1796: Comment on the application submitted by

Dr. W. Lo Petts. Bul. Zool. Nomen. 6: 207-200:

vo.

34.

a -

36.

Oi.

58.

60.

Oi,

G2.

63.

64.

65.

66.

67.

68.

69.

Hee

1953.

1956.

ee cow ge

1957.

Bre

1oS7.,

I958:

(958.

1956.

1958.

(oo.

D9.

LOU.

[S5o.

[960.

|960.

I961.

racl,

Henry K. Townes 11

(with Jose Morales). Control of field rats in the Philippines, with

special reference to Cotabato. Plant Industry Digest, Manila.

Dec. 1953, pp. 3-I2.

The Nearctic species of trigonalid wasps. Proc. U.S. Natl. Mus.

106: 295-304.

The species of Plectochorus (Hymenoptera, Ichneumonidae).

Philippine Jour. Sci. 65: 257-261.

A revision of the genera of Poementiini and Xoridini (Hymenoptera

Ichneumonidae). Proc. Ent. Soc. Washington 59: 15-23.

Nearctic wasps of the subfamilies Pepsinae and Ceropalinae. U.S.

Natl. Mus. Bull. 209, 286 pp.

A review of the generic names proposed for Old World ichneumon-

ids, the types of whose genotypes are in Japan, Formosa, or North

American (Hymenoptera, Ichneumonidae). Proc. Ent. Soc.

Washington 59: 100-120.

A bibliography of the scientific publications of R. A. Cushman.

Proce Enty Soc. Washington $9: 248-254,

Miscellaneous comments on the draft of the "Regles'. Bull. Zool.

Nomenclature !5: 914-930.

The name Psammochares versus Pompilus. Syst. Zool. 6: I5I-I56.

Insects of Micronesia, Hymenoptera: Ichneumonidae, Stephanidae,

and Evaniidae. In Insects of Micronesia 19(2): 35-87.

The application of the name Plectiscus (Hymenoptera, Ichneumon-

idae). Proc. Ent. Soc. Washington 60: 221.

Some biological characteristics of the Ichneumonidae (Hymenop-

tera) in relation to biological control. Jour. Econ. Ent. 5!: 650-

652.

(with M. Townes). Ichneumon-flies of America north of Mexico:

|. Subfamily Metopiinae. U.S. Natl. Mus. Bull. 216(1), 318 pp.

The present condition of the Gravenhorst collection of !chneumon-

idae (Hymenoptera). Proc. Ent. Soc. Washington 61: 76-78.

Notes on the types of Nearctic Tendipedini in London and

Copenhagen (Diptera, Tendipedidae). Proc. Ent. Soc. Washington

Gl? 135.

(with M. Townes, G. S. Walley, et aZ.). Ichneumon-flies of

America north of Mexico: 2. Subfamilies Ephialtinae, Xoridinae,

and Acaenitinae. U.S. Natl. Mus. Bull. 216(2), 676 pp.

The application of the name Syene (Hymenoptera, Ichneumonidae).

Proc. Ent. Soc. Washington 62: 43.

Annotated list of the types of Nearctic ichneumonids in European

museums (Hymenoptera). Proc. Ent. Soc. Washington 63: 103-113.

Some ichneumonid types in European museums that were described

from no locality or from incorrect localities (Hymenoptera).

Proc. Ent. Soc. Washington 63: 165-178.

Pan

Ta.

73.

74,

1D.

76.

(eck

13,

80.

ot.

Sz.

83.

84.

85.

86.

Ohie

88.

89.

I96l.

[96L.

1962.

hgGz,

1962.

1963.

1964.

[965.

1965.

i965.

[e65..

Contrib. Amer. Ent. Inst., vol. 20, 1983

(with M. Townes and V. K. Gupta). A catalogue and reclassifica-

tion of the Indo-Australian Ichneumonidae. Mem. Amer. Ent. Inst.

l, S22 pp.

Some species described as ichneumonids but belonging to other

families (Hymenoptera). Proc. Ent. Soc. Washington 63: 287-289.

(with M. Townes). Ichneumon-flies of America north of Mexico:

3. Subfamily Gelinae, tribe Mesostenini. U.S. Natl. Mus. Bull.

216(3), 602 pp.

A new generic name in the polysphinctine ichneumonids (Hymenop-

tera)... Proc. Ent. Soc. Washington 64; 38.

The synonymy of Zimmeria with Cotiheresiarches (Hymenoptera,

Ichneumonidae). Proc. Ent. Soc. Washington 64: II6.

Host selection patterns in some Nearctic ichneumonids (Hymenop-

tera). Verh. X!I Intern. Kong. Ent. Wien 2: 738-74l.

(with V. K. Gupta). Ichneumon-flies of America north of Mexico:

4. Subfamily Gelinae, tribe Hemigasterini. Mem. Amer. Ent.

Inst. 2, 305 pp.

Design for a Malaise trap. Proc. Ent. Soc. Washington 64: 253-

262.

A revision of Demopheles (Hymenoptera, Ichneumonidae). Proc.

Ent. Soc. Washington 65: 47-50.

Notes on the types of Nearctic Pepsinae and Ceropalinae that are

in London, Lund, Ottawa, and Quebec (Hymenoptera, Psam-

mocharidae). Proc. Ent. Soc. Washington. 65: 15.

Sachtlebenia, a new genus of glyptine ichneumonid (Hymenoptera).

Beitr. Ent. 135262525.

Evidence among some Nearctic Ichneumonidae for a Bering migra-

tion route, pp. 153-158. In Gressitt, Pacific Basin biogeography,

Bishop Museum, Honolulu.

Mislabeled figures in Seyrig's Gelinae of Madagascar. Proc. Ent.

Soc. Washington 66: 18.

Anhang [zu Ubersicht und Bestimmungstabelle der palaearktischen

Gattungen der ehemaligen Unterfamilie Pimplinae auct.]. Ent.

abhand!l. Mus. Tierkunde Dresden 29: 578-579.

Insects of Campbell Island. Hymenoptera: Ichneumonidae.

Pacific Ins. Monog. 7: 496-500.

A new Cremastus (Ichneumonidae), parasitic on Oberea. Papers

Michigan Acad. Sci., Arts, Letters 50: 105-I06.

(with S. Momoi and M. Townes). A catalogue and reclassification

of the eastern Palearctic Ichneumonidae. Mem. Amer. Ent. Inst.

5) Opp.

Labeling in the Gravenhorst collection of Ichneumonidae (Hymenop-

tera). Polskie Pismo Ent. 35: 403-407.

Nomenclatural notes on European Ichneumonidae (Hymenoptera).

Polskie Pismo Ent. 35: 409-417.

90.

a2.

93.

94.

95.

96.

a i

98.

99.

100.

TOL:

102.

103.

104.

106.

107.

108.

109.

110.

1966.

1966

1967.

1967

1967.

i9Oo:

1969.

i969.

196%.

1970.

197 Dy

1970.

L970;

Lore:

l971.

1971,

72.

OTe.

i972.

Henry K. Townes } 13

Notes on three fossil genera of Ichneumonidae (Hymenoptera).

Proc. Ent. Soc. Washington 68: 132-135.

Two ichneumonids described incorrectly as from South America

(Hymenoptera). Proc. Ent. Soc. Washington 68: I80-I8I.

(with M. Townes). A catalogue and reclassification of the Neo-

tropic Ichneumonidae. Mem. Amer. Ent. Inst. 8, 367 pp.

Taxonomic notes on Kentrotryphon, Ktenostilpnus, and

Parentypoma (Hymenoptera, Ichneumonidae). Proc. Ent. Soc.

Washington 69: 58-59.

Notes on the Motschoulsky types of Ichneumonidae (Hymenoptera).

Proc. Ent. Soc. Washington 69: 93.

[Description of Nanium, new genus. Hymenoptera, Ichneumonidae. ]

U.S. Dept. Agr., Agr. Monog. 2, suppl. 2: 174-175.

A new Ateleute from the United States (Hymenoptera: Ichneumon-

idae). Proc. Ent. Soc. Washington 69: 181-182.

The genera of Ichneumonidae, part | (Ephialtinae to Agriotypinae).

Mem. Amer. Ent. Inst. Il, 300 pp.

The Neotropic species of Xanthopimpla (Hymenoptera: Ichneumon-

idae). Proc. Ent. Soc. Washington 71: 82-88.

Zoogeography of the ichneumonid genera with only one or two

species in Europe (Hymenoptera). Polskie Pismo Ent. 29: 347-354.

(with J. Oehlke). Schmiedeknechts Ichneumonidentypen aus der

Kollektion des Museums Rudolstadt. Beitr. Ent. 19: 395-412.

The genera of Ichneumonidae, part 2 (Gelinae). Mem. Amer. Ent.

inst. 12, 6a hop,

The genera of Ichneumonidae, part 3 (Lycorininae to Porizontinae).

Mem. Amer. Ent. Inst. 13, 307 pp.

Tendipes plumosus (Diptera: Tendipedidae) at Solberg Lake,

Wisconsin. Michigan Ent. 2: 82.

(with Shui-chen Chiu). The Indo-Australian species of

Xanthopimpla (Ichneumonidae). Mem. Amer. Ent. Inst. 14, 372 pp.

A review of the Ichneumonidae described by Girault (Hymenop-

tera). Proc. Ent. Soc. Washington 72: 458-470.

The genera of Ichneumonidae, part 4 (Cremastinae to Diplaxontinae)

Mem. Amer. Ent. Inst. 17, 372 pp.

The application of the name Neopimpla (Hymenoptera: Ichneumoni-

dae). Proc. Ent. Soc. Washington 73: 445.

The Ichneumonidae as biological control agents. Proc. Tall Timbers

Conference on Animal Control by Habitat Management 3: 235-248.

(with T. R. Torgersen). Diagnostic features of the subfamilies of

Ichneumonidae. Separate publication. 3 sheets.

The function of the eye stalks in Diopsidae (Diptera). Proc. Ent.

Soc. Washington 74: 85-86.

144,

iar

Tse

114.

115.

116.

dias

118.

149.

120.

121.

te2-

123.

124.

120:

126.

127:

t2os

120:

130.

131.

132.

S72.

Ler2:

i972.

O72.

ioi2.

97s.

S73.

197 3.

ots:

igo.

1973.

I9%2.

(O73.

1973.

1975.

OTT.

IS Ly

[S7s..

1981.

1982.

I9S2.

Contrib. Amer. Ent. Inst., vol. 20, 1983

Incorrect usage of the term "new synonymy". Proc. Ent. Soc.

Washington 74: 229.

Rediscovery of Exochus albiceps (Hymenoptera: Ichneumonidae).

Proc. Ent. Soc. Washington 74: 259.

A note on Zonocryptus luctor (Hymenoptera: Ichneumonidae).

Jour. Ent.:Soc. Southern Africa 35:°177-178.

A light-weight Malaise trap. Ent. News 83: 239-247.

The types of Neotropic Ichneumonidae in the Transvaal Museum.

Proc. Ent..Soc.. Washington 74: 310-3)1.

Comments on the design of insect nets. Ent. News 84: I-7.

The moisture problem in cyanide killing bottles. Ent. News 84:

33.

Deep pinning bottoms for freshly pinned specimens. Proc. Ent. Soc.

Washington 75: 124.

A scientifically designed drawer for insect collections. Ent. News

84: 53-6.

A predaceous scarabaeid. Ent. News 84: 62.

Saturniids as bat mimics. Ent. News 84: 72.

Two ichneumonids (Hymenoptera) from the early Cretaceous. Proc.

Ent. Soc. Washington 75: 216-219.

(with M. C. Townes). A catalogue and reclassification of the

Ethiopian Ichneumonidae. Mem. Amer. Ent. Inst. 19, 416 pp.

Three tryphonine ichneumonids from Cretaceous amber (Hymenop-

tera). Proc. Ent. Soe. Washington 75: 282-287.

The type of Sphex viatica Linnaeus (Hymenoptera, Sphecidae).

Polskie Pismo Ent. 43: 91-96.

The parasitic Hymenoptera with the longest ovipositors with

descriptions of two new Ichneumonidae. Ent. News 86: 123-127.

A revision of the Rhopalosomatidae (Hymenoptera). Contrib. Amer.

Ent: inst..150)), 34app. ;

A revision of the Heloridae (Hymenoptera). Contrib. Amer. Ent.

inst. 15(2 12 pp:

(with M. C. Townes). Ichneumon-flies of America north of Mexico:

7. Subfamily Banchinae, tribes Lissonotini and Banchini. Mem.

Amer. Ent.. inst..26, 614. pp.

(with M. C. Townes). A revision of the Serphidae (Hymenoptera).

Mem. Amer. Ent. Inst. 32, 541 pp.

(with M. C. Townes). A revision of the Serphidae (Hymenoptera).

Mem. Amer. Ent. Inst. 32, 541 pp.

(with M. C. Townes). A description of the Townes collection of

Hymenoptera. !2 pp. Privately printed.

A DESCRIPTION OF THE TOWNES COLLECTION OF HYMENOPTERA

Henry and Marjorie Townes

9950 Warren Road, Ann Arbor, Michigan 48105

The family of Henry and Marjorie Townes has a collection of insects that is

increasingly used in research on Hymenoptera. This is a report on the collec-

tion, to provide information on the material it contains.

The entire collection contains approximately 793, 000 specimens of insects,

pinned and labeled. There is almost no material in alcohol, and only about

3,000 microscope slides, The slides preserve specimens parts, such as geni-

talia and wings, mounted for detailed study, not independent complete speci -

mens, In the collection there are 447, 371 Ichneumonidae, 60,112 Anomalidae

(= Braconidae), and 135, 851 other Hymenoptera. Outside of the Hymenoptera

there are an estimated 150, 000 specimens of other insect orders, including a

reasonably good collection of Neuroptera and Mecoptera, a fair collection of

Trichoptera (pinned), and the midges of the tribe Tendipedini that were used

for H. Townes’ revision of the Nearctic species of that tribe (1945. Amer.

Midland Nat. 34: 1-206). The holdings of Diptera, Coleoptera, Lepidoptera,

Hemiptera, and Homoptera, though respectable for a general reference collec-

tion, are of minor importance for research. The Ephemerida, Odonota, Or-

thoptera, Dermaptera, and Plecoptera were given away in the years 1950 to

1956 and few specimens of those orders have accumulated since then.

In summary, the Townes Collection is now about 81% Hymenoptera and

56% Ichneumonidae.

The collection is located at 5950 Warren Road, Ann Arbor, Michigan, ina

building constructed especially for it. It is contained in 150 cabinets of the

style described by H. Townes in 1951 (Coleopterists’ Bul. 5: 21-27). Each

cabinet holds thirty boxes, 9 x 13 x 2 1/2 inches, of which about half are

wooden Schmidt boxes, the rest cardboard boxes of the same size. The build-

ing also houses a library on insect taxonomy (especially Hymenoptera) and

space and facilities for five taxonomists. The collection has always been in

private ownership, with costs of housing and the addition of specimens borne

by the owners.

The collection was started before 1930 at Greenville, South Carolina. By

the end of 1933 it contained about 4,000 specimens from Greenville Co. and

Horry Co, in South Carolina, from Transylvania Co., North Carolina and from

a few other localities, It was strongest in Lepidoptera, Tipulidae, and

Odonata,

In 1934, the collecting emphasis shifted to the Ichneumonidae and has

remained there since, with secondary emphasis on other parasitic Hymenop-

tera and on sawflies and wasps.

Published as a separate pamphlet, February, 1982.

16 Contrib. Amer. Ent. Inst., vol. 20, 1983

HYMENOPTERA CONTENT OF COLLECTION

The number of specimens of each family of Hymenoptera are tabulated

below. For this list the taxonomic limits of families and the family arrange-

ment correspond to those in Krombein, et al., 1979, ‘ ‘Catalog of Hymenop-

tera in America North of Mexico.’’ Although the taxonomy of the Catalog is

used, it is not necessarily endorsed, The names of the families in the list

below correspond with the usage in the Townes Collection, with the equiva-

lents used in the Catalog given in parentheses,

Hymenoptera specimens in the Townes Collection by family,

as of December 1981

Megalodontoidea 556

Xyelidae 253

Pamphiliidae 303

Tenthredinoidea 13, 339

Pterygophoridae 1,157

(= Pergidae)

Cryptidae 1,038

(= Argidae)

Clavellariidae 126

(= Cimbicidae)

Pteronidae 31

(= Diprionidae)

Tenthredinidae 10, 987

Siricoidea 268

Xiphydriidae 65

Siricidae 91

Orussidae 112

Cephoidea 619

Cephidae 619

Ichneumonoidea 509, 557

Apozygidae 1

Anomalidae 60,112

(= Braconidae)

Incubidae 1, 690

(= Aphidiidae)

Hybrizontidae 67

Ichneumonidae 447, 391

Stephanidae 288

Megalyridae 28

Chalcidoidea 13, 239

Callimomidae 1, 340

(= Torymidae)

Pteromalidae 3, 745

Eurytomidae 939

Chalcididae 3, 828

Leucospidae 129

Eucharitidae 507

Eupelmatidae 426

Encyrtidae 274

Eulophidae 1, 755

Mymaridae 296

Cynipoidea 4, 086

Ibaliidae 84

Liopteridae 15

Figitidae 923

Eucoilidae 2, 399

Alloxystidae 124

Cynipidae 591

Evanioidea 11, 339

Evaniidae 7, 958

Aulacidae 1,275

Gasteruptiidae 2, 106

Pelecinoidea 255

Pelecinidae 255

Serphoidea 15, 020

Roproniidae 86

Austroniidae 3

Monomachidae 375

New family 1

Heloridae 142

Serphidae 4, 201

Diapriidae 7, 583

Sparasionidae 2, 081

(= Scelionidae)

Platygastridae 548

Ceraphronoidea 275

Ceraphronidae 71

Megaspilidae 204

Trigonaloidea 340

Trigonalidae 340

Chrysidoidea 12, 381

Plumariidae 190

Bethylidae 5, 928

The Townes Collection of Hymenoptera 17

Clystopsenellidae 19

(= Scolebythidae)

Sclerogibbidae 118

Chrysididae 2, 354

Dryinidae 3, 613

Embolemidae 159

Scolioidea 15, 527

Tiphiidae 9, 574

Sierolomorphidae 149

Mutillidae 4, 802

Scoliidae 795

Sapygidae 207

Formicoidea 1, 894

Formicidae 1, 894

Vespoidea 5, 998

Masaridae 107

Eumenidae 1, 933

Vespidae 3, 958

Psammocharoidea 20, 183

Psammocharidae 19, 820

(= Pompilidae)

Rhopalosomatidae 355

Sphecoidea 13, 396

Ampulicidae 209

Sphecidae 1, 909

Pemphredonidae 2, 082

Astatidae 117

Larridae 3, 465

Crabronidae 2, 739

Mellinidae 22

Bembicidae 1, 820

(= Nyssonidae)

Philanthidae 1, 033

Apoidea 5, 062

Hylaeidae 827

(= Colletidae)

Andrenidae 397

Halictidae 1, 967

Mellitidae 10

Megachilidae 440

Lasiidae 1,033

(= Anthophoridae)

Apidae 388

Total Hymenoptera 643, 334

Since the numbers of Anomalidae and Ichneumonidae are very large, the

holdings of these two families are listed by subfamily. The subfamily limits

(but not always the names) are as in the ‘‘ Catalog of Hymenoptera of America

North of Mexico’’, except that the Stilbopinae are separated from Banchinae

(Ichneumonidae).

Specimens of Anomalidae (= Braconidae) in the Townes Collection,

as of December 1981

Spathiinae 3, 854

(= Doryctinae)

Vipiinae 8, 097

(= Braconinae)

Hormiinae 943

(= Exothecinae)

Rogadinae 6, 214

Meteorideinae lv

Helconinae 1,079

Zelinae 479

Macrocentrinae 2, 346

Amicrocentrinae 4

Xiphozelinae 3

Braconinae 4, 083

(= Agathidinae)

Opiinae 2, 225

Alysiinae 3, 902

Sigalphinae 49

Anomalinae 5, 404

(= Cheloninae)

Adeliinae 12

Microgastrinae 13, 002

New subfamily 16

Ichneutinae 542

Blacinae 3, 869

Neoneurinae 21

Leiophroninae 3, 958

(= Euphorinae)

Total Anomalidae 60, 112

Contrib. Amer. Ent. Insts vol 20, 1983

Specimens of Ichneumonidae in the Townes Collection,

Ephialtinae 24, 398

Tryphoninae 24, 445

Eucerotinae 668

Labiinae 1,556

Adelognathinae 529

Xoridinae 1, 684

Agriotypinae 4

Gelinae 110, 103

Lycorininae 275

Stilbopinae 136

Neorhacodinae 7

Banchinae 23, 884

Scolobatinae 23, 459

Porizontinae 70, 048

as of December 1981

Cremastinae 9, 970

Phrudinae 149

Tersilochinae 3, 458

Ophioninae 12, 295

Mesochorinae 7, 744

Metopiinae 10, 085

Anomaloninae 8, 288

Acaenitinae 791

Microleptinae 15, 258

Orthopelmatinae 324

Collyriinae 84

Orthocentrinae 10, 818

Diplazontinae 10,021

Ichneumoninae 176, 890

Total Ichneumonidae 447, 371

GEOGRAPHIC AREAS REPRESENTED

There are notes on the origin of the lots coming into the collection,

especially the specimens of Ichneumonidae, For the ichneumonids one can

compile a list of the countries of origin of most of the specimens, and the

origins of the ichneumonids are a good indication of the origins of the specimens

of other families, which were generally collected at the same times and places.

In many cases the count of ichneumonids per country is precise, in others the

number is given as a round number (estimated). The tabulations from different

localities is inexact because numerous small lots were not accessioned, be-

cause of attrition by gifts, and the increased variety of locations resulting from

exchanges, The tabulation below totals only 439, 623 ichneumonids while a count

of the collection shows it to contain 447, 371 (plus 1, 639 holotypes).

Origins of the specimens of Ichneumonidae

North America (Ichneumonidae per country)

Canada 16, 860

Costa Rica 6, 646

Greenland 50

Guatamala 20

Mexico 3, 100

Panama 347

United States 188, 861

West Indies 445

North America total 216, 359

The Townes Collection of Hymenoptera

South America (Ichneumonidae per country)

Argentina 10, 400 Paraguay 257

Bolivia 1, 048 Peru 4,274

Brazil 36, 264 Surinam 1,073

Chile 4,810 Trinidad 679

Colombia 938 Venezuela 4,990

Ecuador 0, 900

south America total 10, 633

Europe (Ichneumonidae per country)

Austria 2,676 Iceland 20

Bulgaria 25 Ireland 000

Cyprus 1, 340 Italy 6, 645

Czechoslovakia 1,053 Poland 20

Denmark 1, 000 Russia 1, 707

East Germany 1, 000 Scotland 399

England 16, 504 Spain 030

France 329 Sweden 16, 551

Finland 1,000 Switzerland 83

Hungary 150 West Germany _ 3, 500

Europe total 00, 111

Asia (Ichneumonidae per country)

Burma 400 Nepal D2

China 2,927 Siberia 150

India 7, 466 South Korea 444

Japan 9, 786

Asia total 21, 225

Africa (Ichneumonidae per country)

Angola 200 Rhodesia 390

Botswana 42 Sierra Leone 2, 990

Ghana 30 South Africa 22, 286

Kenya 3, 696 Swaziland 25

Madagascar 4,075 Tanzania 1,180

Mozambique 20 Uganda 3, 000

Nigeria 1, 316 Zaire 826

Africa total 40, 406

Contrib. Amer. Ent. Inst., vol. 20, 1983

Pacific Area (Ichneumonidae per country)

Australia 12, 769

Indonesia 1, 700

Micronesia. 30

New Caledonia 20

New Guinea 6, 877

New Zealand 1,016

Philippines 12, 500

Solomons 150

Taiwan 826

Pacific Area total 35, 938

Total tabulated, all countries 439, 623

SOURCES OF SPECIMENS

Approximately 55% of the specimens in the collection were collected by the

owners, The rest were received from other sources by purchase, as gifts, in

exchange, or as payment for taxonomic work. Most specimens were received

unmounted in papers or in alcohol. After unwanted duplicates were culled from

the lots, the remaining specimens were mounted and labeled by the owners.

The localities in which specimens were collected by the owners themselves are

spotted on an accompanying map. A second map shows the origins of other

Specimen lots. Localities for minor sources are not indicated.

George Townes helped Henry and Marjorie Townes collect in the western

United States in 1948. Jean Townes and David Townes helped collect in the

western United States in 1947 and 1948, in the Philippines 1952-1954, and in

Japan in 1954, Chloe West assisted M. Townes on a collecting trip to Europe

in 1964, Specimens collected by M. Townes 1935-1937 are labeled with her

maiden name (M. Chapman). The other collectors who contributed noteworthy

lots are listed below,

Collectors Making Substantial Additions to the Collection,

and their Localities

Moacir Alvarenga: Brazil

Pablo Anduze: Venezuela

Ross Arnett: Florida

P. P. Babiy: New York

John Bain: New Zealand

Ian Bampton: Kenya

Bryan Beirne: Ireland

Jackie Belwood: Ontario

Fred Bianchi: India

G. E. Bohart: Colombia & Mexico

R. M. Bohart: California & Ryukyus

M. S. Bourgeois: Zaire

Sam Breland: Kentucky & Panama

Nancy Breisch: Massachusetts

T. Cekalovik: Chile

Shui-chen Chiu: Taiwan

William Clarke-MacIntyre: Ecuador

C. R. Cunningham-van Someren:

Kenya

R. & K. Dreisbach: Mexico &

Michigan

K. M. Fender: Oregon

H. R. Foxlee: British Columbia

A. E. Freeman: Ellesmere Island

Franco Frilli: Italy

Manfredo Fritz: Argentina & Bolivia

David Fullaway: Hawaii

D. C. Geijskes: Surinam

Fred Gess: South Africa

Peter Ginn: Botswana & Zimbabwe

J. L. Gressitt: China, Japan, &

Taiwan

V. K. Gupta: India

Erasmus Haeselbarth: South Africa

The Townes Collection

Bernd Heinrich: New Guinea

Gerd Heinrich: Angola, Austria,

Burma, South Africa, Tanzania,

Uganda, and West Germany

Henry & Anne Howden: Australia

Colombia, Costa Rica,

El Salvador, Ecuador, Honduras,

Mexico, Panama, & Trinidad

Rex Jubb: South Africa

Dan Janzen: Costa Rica

Karl Krombein: Solomons & West

Virginia

C. Lamberton: Madagascar

G. A. Lancaster: Uganda

M. A. Lieftinck: Indonesia

J. Lima: Brazil

Luis Ling: Ecuador & Ontario

Linda Losito: England

T. C. Maa: China

Jan Maéek: Czechoslovakia

W. R. M. Mason: Costa Rica &

Ecuador

Lubomir Masner: Dominican

Republic

G. A. Mavroumoustakis: Cyprus

John Medler: Nigeria

Karl Miller: Sweden

P. S. Nathan: India

I, Nikitin: Australia

D. Novicky: Austria

S. J. Oliveira: Brazil

Denis Owen: England, Sierra Leone,

& Uganda

Jennifer Owen: England

Agoroacheri Owiny: Kenya

Reginald Painter: Mexico

Charles Palmer: Costa Rica

Stuart & Jarmila Peck: Japan,

New Mexico, & Texas

of Hymenoptera

Luis Pena: Argentina, Bolivia,

Chile, Ecuador, Paraguay,

& Peru

Fritz Plaumann: Brazil

Charles Porter: Argentina

O. Ranin: Finland

Richard Reardon: New Jersey &

Pennsylvania

C. L. Remington: New Caledonia

J. G. Rempel: Saskatchewan

C. & M. Rettenmeyer: Panama

Peter Rush: Alaska, Louisiana, &

Michigan

Kaku Sato: Japan & Korea

J. Sedlacek & J. H. Sedlacek:

New Guinea & Australia

Charles Seydel: Zaire

A. Seyrig: Madagascar

Peter Shanahan: New Guinea

Lionel Stange: Argentina

J. Stibick: New Britain, New

Guinea, & Tasmania

B. R. Stuckenberg: South Africa

Bo Svenson: Sweden

Charles Townes: China & Nepal

David Townes: Alaska, Denmark,

Italy, Sweden, & West Germany

George & Laura Townes:

South Carolina

Jean Townes: Nigeria & Oregon

C. A. Triplehorn: Panama

M. E. Walsh: Java

F. H. Walz: Bolivia

A. M. R. Wegener: Moluccas

Mary Jane West-Eberhard: Colombia

Vincent Whitehead: South Africa

Felix Woytkowski: Peru

From 1938 to 1955, collections for naming were regularly received from

individuals and universities, with an agreement that payment for the taxonomic

work would be a selected portion of the specimens. These lots furnished a

valued miscellany of minor additions to the collection. Some of the most impor-

tant lots were from the collections in Columbus, Ohio; Edmonton, Alberta;

Bogor, Indonesia; and the Dreisbach Collection.

| Major exchanges of Ichneumonidae were transacted with the museums in

Washington, Sapporo, London, Copenhagen, and East Berlin, and with the pri-

vate collections of V. K. Gupta, Gerd Heinrich, H. G. M. Teunissen, and A.

Roman, There were smaller exchanges with the museums in Ottawa, Paris,

Leningrad, and Moscow, and with the private collections of J. Aubert and M.

Constantineanu.,

Paratypes were included in most of these exchanges.

1983

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ARRANGEMENT OF SPECIMENS

The arrangement of the collection is taxonomic, with only a little segrega-

tion by geographic regions. All specimens are sorted to family, and in the

Ichneumonidae and Anomalidae to subfamily. In certain sections, especially

in the Ichneumonidae, the unstudied specimens are sorted to genera, Where

Specimens are named to species, a set of specimens (5 males and 6 females

of each species) is kept in a main reference collection and any additional

Specimens are put in a ‘‘duplicate’’ collection. Thus, the main reference

collection is synoptic, and the long series are in the ‘‘ duplicate’’ collection,

Specimens not named to species are grouped at the end of each genus, tribe,

subfamily, or family, depending on how precisely they have been sorted.

HOLOTYPE SPECIMENS

Holotype specimens are kept separately from the general collection,

arranged in sequence by a number assigned to each, There is a card file of

the type names, giving the number under which each can be found, reference

to the place of publication, and collecting data.

The numbers of types are listed below, by family and byauthor. This list

does not include manuscript types nor types in the hands of the authors who

have not yet returned them to the Townes Collection,

_ List of the Holotypes in each Family

Danaidae (Lepidoptera): Danaus davidi Schréder.

Tendipedidae (Diptera): 63 types of Tendipedini by H. Townes.

Bibionidae (Diptera): Bibio townesi Hardy.

Anomalidae (= Braconidae). This and all of the following families are

Hymenoptera,

374 types, by C. van Achterberg (13), Clare Baltazar (2), Shama Bhat (2),

Shama Bhat and V. K. Gupta (24), Max Fischer (318), John Martin

(Aliolus denticulatus), W. R. M. Mason (Muesebeckia chota), Gilbert

Nixon (12), and Garland Riegel (Alysia townesi).

Ichneumonidae. 1,651 types, by Douglas Allen (3), Jacques Aubert (6), Clare

Baltazar (47), John Barron (14), B. D. Burks (2), R. W. Carlson

(Triclistus minutus), Girish Chandra and V. K. Gupta (24), Shui-chen Chiu

(Yezoceryx townesi), R. A. Cushman (Aulophion excarinatus), Clement

Dasch (450), Alex Delobel (2), Ian Gauld (6), Ian Gauld and Pam Mitchell

(78), David Grimble (Barylypa irona), M. L. Gupta (5), V. K. Gupta (23)

V. K. Gupta and Joseph Jonathon (6), V. K. Gupta and Sharda Maheshwary

(15), V. K. Gupta and D, T. Tikar (22), Gerd Heinrich (157), Rolf Hinz (3),

Klaus Horstmann (6), Keith Johnson (Xiphosomella setoni), J. K. Jonathan

(11), M. K. Kamath and V. K. Gupta (20), D. Kasparyan (8), Patrick

Kennedy (Mnioes lunatus), G. J. Kerrich (Anisoctenion clauseni), Charles

Krebs (Phytodietus rutilus), Raminder Kaur and Joseph Jonathon (9),

W. R. M. Mason (22), R. T. Mitchell (Acroricnus stylator townesi),

Setsuya Momoi (5), Charles Porter (63), Harry Pratt (Cryptus crassulus),

G. van Rossem (7), John Schmid (3), C. N. Slobodchikoff (2), Hewson Swift

(Hoplismenus teres), Timothy Tigner (3), Torolf Torgersen (5), Toichi

The Townes Collection of Hymenoptera

Uchida (8), Henry Townes (412), Henry and Marjorie Townes (103),

Henry Townes and Shui-chen Chiu (48), Henry Townes and V. K. Gupta

(30), and G. S. Walley (12).

Gasteruptiidae. 4 types by H. Townes.

Roproniidae. Ropronia townesi Yasumatsu.

_Serphidae, 162 types, by H. Townes.

Sparasionidae (= Scelionidae). 5 types, by Lubomir Masner (2) andC. F. W.

Muesebeck (3),

Bethylidae, 8 types, by H. E. Evans.

Tiphiidae. Tiphia tenuis Allen,

Sierolomorphidae, 2 types, by H. E. Evans.

Eumenidae, 2 types, by J. van der Vecht,

Vespidae, 4types, by J. van der Vecht.

Rhopalosomatidae, 9 types, by H. Townes.

Psammocharidae, 18 types, by R. R. Dreisbach (3), H. E, Evans (7),

Henry Townes (5), and Raymond Wahis (3).

Pemphredonidae, 23 types, by J. P. van Lith.

Larridae, Nitela townesorum Krombein.

Crabronidae, 17 types, by Jean Leclercq.

Hylaeidae, Hylaeus packardi Mitchell.

Lasiidae. Nomada townesi Mitchell.

Total number of types: 2,275.

The following ichneumonid types by Gauld and Mitchell were published

erroneously as belonging to the Townes Collection. They belong actually to

the Leiden Museum: Enicospilus amasus, E. arenus, E. fallax, E, isolde,

. laridus, E. olthopi, E, paniscus, E. sambucus, E. serphus, E, spathius,

. toxopeus, E.vernalis, E. vespus, E. zemiotes, Leptophion magus and

nodus.,

The following ichneumonid types by Townes, by Townes and Townes, or

by G. S. Walley were published as belonging to the Washington Museum but

are actually the property of the Townes Collection. (The reason for the

incorrect publication was a misunderstanding by the Smithsonian’s editor. )

The types are listed below by subfamily, in the page order of publication.

METOPIINAE: Chorinaeus longicalcar suturalis, C. aequalis, C. funebris

clarus, C. recurvus, C. labiosus, C. emorsus, Trieces tegularis, T.

sapineus litus, T. ejectus, T. aquilus, T, dentatus, Metopius krombeini

epixanthus, M. vittatus, M. scapulatus, M. errantius californicus, Triclistus

occidentis, T. rectus, Colpotrochia fultoni, Bothromus minoris cruralis,

Carria dreisbachi californica, C. inculcata, Macromalon montanum,

Hypsicera fulviceps, H. cuneata cuneata, Exochus elimatus, E. montivagus,

E. ochreatus, E, spinalis, E, atriceps atricornis, E. sulcatus, E, tenebro-

sus, E. armillosus, E. cnemidotus, E, capnodes, E, canidens, E, quadradens,

E. mesodon, and E, ventricosus. EPHIALTINAE: Anastelgis terminalis,

Zaglyptus arizonicus, Laufeia navajo, Acrodactyla jubata, A. ocellata,

Scambus arizonensis, S. subtilis, S. tenebrosus, Eruga atrata, Piogaster

maculata, Zatypota patellata, Z. favosa, Z. exilis, Z. luteipes, Z. crassipes,

Itoplectis fustiger, and Coccygomimus stricklandi, KORIDINAE: Aplomerus

arugosus, Odontocolon pullum, O. depressum, O. sierrae, O. parvum, O.

dreisbachi, Xorides pictus, and X. semirufus. LABIINAE: Labena grallator

ochreata, ACAENITINAE: Coleocentrus manni coloradensis, GELINAE:

Agrothereutes montanus, A. pallipennis, A. linnae linnae, A, linnae pacifica,

A. albicollaris clypeator, A. notata sierrae, Pycnocryptus alexanderi,

eel el

Contribu Amer. Ent. ‘Inst., vol: 20, 1983

Ischnus laurae, I. velutinus, I. laevifrons, I. politus, Habrocryptoides pictus,

H. lumbarius, Trachysphyrus fasciatus, T. labrator satoi, T. lochmaius, T.

luctuosus atrifemur, T. mentigus mentigus, T, krombeini, T. rugifrons,

Lanugo schlingeri, Compsocryptus unicolor, Joppidium densum, Trychosis

semirubra arizonica, T. kathrynae, T. anagmus, Diapetimorpha brunnea,

Listrognathus bicolor, L. femorata, Cryptohelcostizus nigricans, and

Helcostizus subrectus,

A COMPARISON OF THE TOWNES COLLECTION OF PARASITIC

HYMENOPTERA WITH THAT OF THE SMITHSONIAN INSTITUTION

A comparison of a sizable insect collection with that of the

Smithsonian Institution is of interest because the Smithsonian Collection sur-

passes others in the United States in the same manner that the Library of

Congress surpasses other libraries.

The Townes and the Smithsonian collections can be compared in several

areas ~— size, species and geographic representation, quality of specimens,

curating, and type material. Size (number of specimens) is the only thing

that can be objectively quantified, and that is done below. The table enu-

merates specimens that are mounted, labeled, and sorted at least to family.

Specimens less completely processed are considered not yet available. The

comparisons refer only to the parasitic Hymenoptera as the Townes Collec-

tion has no pretensions elsewhere. Counts were made of the Townes Collec-

tion in December I981, of the Smithsonian Collection in June 1982.

The Smithsonian Collection has been forming for 100 years, receiving

specimens from private individuals, and from a great variety of state and

government agencies. Its specimens are approximately 30% from outside of

the United States, the other 70% mostly from the eastern United States.

Its special strengths are in type specimens and in reared specimens.

The Townes Collection was assembled over a 50 year period, with sub-

stantial help from about 50 contributors. Its main goal was to gather a

representation of the world fauna of certain families of parasitic Hymenop-

tera. The specimens are approximately 60% from outside of the United

States. There are almost no reared specimens. There is a substantial num-

ber of types but not nearly as many as in the collection at the Smithsonian.

In numbers, the Townes Collection is 72% as large as that of the

Smithsonian, as tabulated below. It has a lead in 50 of the families and

subfamilies tabulated and is behind in 36. Its special strength is in the

family Ichneumonidae, in the numbers of foreign specimens, and in its good

condition. Its special weakness is in its representation in the Chalcidoidea.

Although only 72% as large by numbers as the Smithsonian Collection it is

about the same in research value, since about 15% of the Smithsonian Collec-

tion (96,000 of Ichneumonidae and Braconidae) is not sorted well enough to

be accessible and another 15% consists of excessively: long series of minor

use for taxonomy.

The Townes Collection is scheduled to move to Gainesville, Florida where

it will be associated with the Gupta Collection and the Florida State Collec-

tion of Arthropods. These two collections contain an estimated 75,000, and

60,000 specimens of parasitic Hymenoptera respectively. The three collec-

tions combined will total approximately the same as the Smithsonian Collec-

tion.

28 Contrib. Amer. Ent. igt.; Vol. 20; 3 1983

NUMERICAL COMPARISON OF TOWNES AND SMITHSONIAN COLLECTIONS

Townes Smithsonian

Collection Collection

Ichneumonoidea (509, ae (399, 597)

Apozygidae 9

Diecondas (60, 112) (180, 899)

Spathiinae 3, 854 16, 997

Vipiinae 8, 097 21, 088

Hormiinae 943 2, 200

Rogadinae 6, 214 8, 859

Meteorideinae 10 15

Helconinae 1, 079 1,172

Zeleinae | 479 347

Macrocentrinae 2, 346 5, 962

Amacrocentrinae 4 0

Xiphozelinae 3 2

Braconinae 4, 083 we

Opiinae 2, 225 ;

Alysiinae 3. 902 5, 012

Sigalphinae 49 200

Cheloninae 5. 494 9, 962

Adeliinae - 12 72

Microgastrinae _ 13, 002 32, 984

New subfamily 16 0

Ichneutinae 542 302

Blacinae 3, 869 ; 8, 092

Ypsitocerinae 0 O7

Neoneurinae 21 88

Leiophroninae 3, 958 2 soi

Not sorted 0 ?

Aphidiidae 1, 690 11, 164

Hybrizontidae 67 46

Ichneumonidae (447, 391) wr Aon

Ephialtinae 24, 398 ?

Tryphoninae 24, 445 0, 700

Eucerotinae 668 233

Labiinae 1, 556 350

Adelognathinae 529 Oa

Xoridinae 1, 684 478

Agriotypinae 4 12

“peel e an is 102

Stilbopinae 136 11

Neorhacodinae " 8

Banchinae 23, 884 6, 9190

Scolobatinae 23, 459 3, 3854

Porizontinae 70, 048 19, 794

Cremastinae 9, 970 5, 424

Phrudinae 149 27

Townes: Collections 29

Townes Smithsonian

Collection Collection

Tersilochinae 3, 458 1, 643

Ophioninae 12, 295 7, 380

Mesochorinae 7, 744 1, 964

Metopiinae 10, 085 1, 759

Anomaloninae 8, 288 4, 621

Acaenitinae 791 022

Microleptinae 15, 258 765

Orthopelmatinae 324 048

Collyriinae 84 112

Orthocentrinae 10, 818 804

Diplazontinae 10, 021 2,174

Ichneumoninae 76, 890 21, 440

Not sorted 0 o1, 859

Stephanidae 288 497

Megalyridae 28 9

Chalcidoidea (13, 239) (262, 778)

Torymidae 1, 340 22, 073

Pteromalidae 3, 745 09, 659

Eurytomidae 939 63, 258

Chalcididae 3, 828 16, 620

Leucospidae 129 719

Eucharitidae 509 2, 765

Eupelmatidae 426 10, 617

Encyrtidae 274 25, 375

Eulophidae 1, 755 O71, 575

Mymaridae 296 4,019

Cynipoidea (3, 421) (11, 213)

Ibaliidae 84 249

Liopteridae 19 45

Figitidae 923 2, 595

Eucoilidae 2, 399 8, 324

Alloxystidae 124 669

Evanioidea (11, 339) (4, 008)

Evaniidae 1, 958 2, 780

Aulacidae 1, 275 601

Gasteruptiidae 2,106 627

Pelecinoidea (255) (126)

Pelecinidae 255 126

Serphoidea (15, 020) (40, 745)

Roproniidae 86 21

Austroniidae 3 1

Monomachidae 375 30

New family 1 0

Heloridae 142 95

Serphidae 4,201 2, 521

Diapriidae 7, 583 12,115

Scelionidae 2,081 17, 592

Platygastridae 548 8, 370

30 Contrib. Amer. Ent. Imst.; vol. 20,

Townes

Collection

Ceraphronoidea (275)

Ceraphronidae 71

Megaspilidae 204

Trigonaloidea (340)

Trigonalidae 340

Chrysidoidea (10, 024)

(except Chrysididae)

Plumariidae 190

Bethylidae 5, 928

Scolebythidae 19

Sclerogibbidae 118

Dryinidae 3, 613

Embolemidae 159

Scolioidea (10, 518)

(except Mutillidae &

Sapygidae) :

Tiphiidae 9,574

Sierolomorphidae 149

Scoliidae 795

Psammocharoidea (355)

(except Psammocharidae)

Rhopalosomatidae 355

1983

Smithsonian

Collection

(4, 109)

Total parasitic Hymenoptera 561,107

174, 206

THE ARTICLES PRESENTED IN THIS COMMEMORATIVE VOLUME

ARE

INDIVIDUALLY AND: COLLECTIVELY DEDICATED TO

HENRY K. TOWNES

FOR HIS WORK AND LEADERSHIP IN THE STUDY OF THE

ICHNEUMONIDAE AND OTHER PARASITIC HYMENOPTERA

A HOLE IN A TENT OR HOW TO EXPLORE

INSECT ABUNDANCE AND DIVERSITY

Denis F. Owen

Department of Biology, Oxford Polytechnic

Headington, Oxford OX 3 OBP, U.K.

Every scientist likes to think that his contribution will be incorporated

into the text books and that his name will be remembered. But theories and

hypotheses can be tested and shown to be wrong, and data can be reworked

and reinterpreted to show that the researcher missed the point, collected the

wrong information, or performed the wrong experiment. There is, however,

one activity open to the biologist which ensures that his name is recorded

and remembered: discovering a new, undescribed species of plant or animal

and having it named after him. Even if it later turns out the species has al-

ready been described, perhaps in an unlikely journal or in an unfamiliar

language, the name is preserved in the archives of synonymy.

In 1964 and 1965 I collected a sample of Ichneumonidae in a Malaise

trap sited in the garden of the house I lived in at Kampala, Uganda. The

sample was sent to Henry Townes and in it he found 88 males and 41 fe-

males of a genus and species new to science. The genus was described as

Owenus and the species as Owenus minor (Townes 1970). The original drawing

of Owenus minor is reproduced in Fig. |. Further collecting in Africa and

examination of collections of ichneumons in museums has revealed at least

44 species of Owenus, but to date only three have been described.

A specialist in ichneumons would not be astonished to hear of the dis-

covery of a new genus with more than forty species from the heart of

Africa, but an ecologist without special knowledge of ichneumons is at once

impressed and humbled by the extraordinary diversity of the family with

more undescribed than described species in some areas. I am of course

thrilled to have a genus named after me. I may regret that the rules of

nomenclature do not allow the name Owenus to be used again for a more

spectacular animal, a new shark or parrot perhaps, but in the event I am

unlikely to discover these.

The genus Owenus was discovered as a consequence of taking routine

samples of insects in a Malaise trap to try and detect seasonal changes in

abundance and diversity at a site near the equator where seasonal changes

in weather are not marked. The value of the Malaise trap as a means of

assessing the insect fauna of a site is beginning to be appreciated and var-

ious designs of trap are in wide use. I was first introduced to the Malaise

trap in 1960. Henry Townes suggested that it might be interesting to oper-

ate one on Evans Old Field on the Edwin S. George Reserve, Michigan, an

intensively studied site where, it was believed, the insect fauna was well

known. The trap dramatically increased the species list, especially of

Diptera and the smaller Lepidoptera (Evans and Murdoch 1968), and also

demonstrated that various types of quantitative information could be obtained

(Evans and Owen 1965). I have used Malaise traps ever since 1960 and in

this paper I shall review the kinds of ecological information that can be ob-

tained from them. A partial bibliography of the Malaise trap is given by

Steyskal (1981). This is valuable but it does not indicate how the trap can

be used as an effective sampling device.

34 Contrib... Amer. tnt. Inst., vol. 20, 1983

Fig. 1. Owenus minor Townes

A SIMPLE IDEA

There is no doubt that Rene Malaise discovered what has since become

known as the Malaise trap. In his account (Malaise 1937) of a new insect

trap he wrote, "During my extensive travels I have repeatedly found that in-

sects happened to enter my tent, and that they always accumulated at the

ceiling-corners in vain efforts to escape at that place without paying any at-

tention to the open tent-door. On one occasion one of the upper tent-

corners happened to have a small hole torn in the fabric, and through this

hole all the insects pressed their way and escaped. Later the idea occurred

to me, that, if insects could enter a tent and not find their way out, and

always persistently tried to reach the ceiling, a trap made as invisible as

possible and put up at a place where insects are wont to patrol back and

forth, might catch them much better than any tent and perhaps better than

a man with a net, as a trap could catch all the time, by night as well as

by day, and never be forced to quit catching when it was best because dinner-

time was at hand." Malaise then described how he used the new device in

Burma and what an extraordinary number and variety of insects it produced.

He also made a prediction and wrote, "For students interested in a statistic-

al survey of the insect-fauna of a particular region, this trap may prove most

useful, and it is possible that it might also be used with success in orchards

to decimate insect-pests."" Despite Malaise's enthusiasm for his discovery,

the idea was not taken up by many entomologists until Henry Townes, with

customary precision and attention to detail, explained exactly how to make

a trap (Townes 1962). This particular design, open on four sides, was the

D. Owen: A hole in a tent 35

one I used in Michigan and later in Uganda. It was rather time-consuming

to construct and soon after a simpler type, open on two sides, was invented

(Townes 1972a), I adopted it and have used it ever since.

A suitably-sited Malaise trap catches an extraordinary number and

variety of insects. Since no attractant is used, the trap samplés only those

individuals that enter its air space and thus accurately monitors what is go-

ing on in that space. If a trap is left at the same site for a whole season

an enormous amount of information is obtained. It is possible to compare

sites and to compare seasons, but impossible to do the necessary taxonomic

work on all the insects collected. The trap that caught the 129 specimens

of Owenus minor in the Kampala garden caught in one year an estimated

113,538 insects belonging to 13 orders. The insects were not counted out

individually and the estimate was obtained by counting the insects in 52

48-hour samples taken once a week throughout the year. I have never at-

tempted to estimate the number of insects taken in a year in a Malaise trap

operated in a temperate climate. I suspect the figure is just as high with,

of course, much more conspicuous seasonal changes in numbers.

Rene Malaise's observation of what insects did when they reached the

hole in the top of his tent resulted in a significant break-through in the

sampling of flying insects. Henry and Marjorie Townes have together been

responsible for what might be called research and development in the con-

struction and operation of Malaise traps; indeed there was a time when the

traps almost became known as Townes traps.

Nearly everywhere I have used Malaise traps I have taken out the

Ichneumonidae and sent them to the American Entomological Institute. My

specimens are always collected and preserved in 70% alcohol and Marjorie

Townes has pinned and dried them. Henry Townes then tabulated the

species. One result of this collaboration is that the American Entomological

Institute received some interesting and valuable specimens; another is that

the tabulations of numbers and species have raised some questions about com-

parisons of temperate and tropical species diversity.

ICHNEUMONID DIVERSITY AT FOUR SITES

Ichneumonids tend to occur as individuals. Rarely is a species common

and samples taken in Malaise traps invariably contain many species. With

an estimated 60,000 species in the world, more than all the vertebrates put

together (Townes 1972b), they are an excellent group for comparisons of

species diversity at different sites. Numerical information on diversity is

available from four samples, two tropical and two temperate. The first sam-

ple obtained was from the garden at Kampala, Uganda, and the results were

published in Owen and Chanter (1970). Next came a similar sample from a

garden at Freetown, Sierra Leone (Owen 1971). Then followed two temperate

samples, one from a garden in Leicester, England (Owen, Townes and Townes

1981), the other from disturbed land along a small stream in Skane, Sweden

(Owen and Owen 1974). The four sites are not of course identical. Kampala

and Freetown are at opposite sides of tropical Africa, the garden at

Leicester is decidedly suburban, while the Sk&ne site is not a garden at all.

But each was obtained by using Malaise traps for 12-month periods, and

36 Contrib. Amer. Ent. Inst., vol. 20, 1983

the only assumption made is that the traps used did not select for either

rare Of common species.

The four samples are summarized in Table 1. The information theory

index of diversity, H, enables comparison of species diversity. The values

of H for Freetown and Leicester are not significantly different but all the

other paired samples are different from one another at the 1% level. Par-

ticularly interesting is the fact that H is significantly smaller for the

Kampala sample than for the Leicester and Sk&ne samples, and is smaller

for the Freetown sample than for the Sk&ne sample.

TABLE 1. Two tropical and two temperate samples of Ichneumonidae com-

pared. From Owen and Owen (1974).

S Diversity Species %

eae Species index, H, taken commonest

sa and s.e. once species

Kampala 2,268 293 4.524 + 0.032 116 10.1

Freetown 1,979 319 4.934 + 0.029 117 4.9

Leicester 2,495 326 4.937 + 0.024 27 402

+ 0.014 203 345

Skane 10,994 758 5.481

In Table | no special attention should be paid to the actual sample

sizes nor to the number of species in each sample, except to note that at

all four sites the number of species is very high indeed. More relevant is

that at all sites the number of species taken once only is consistently high

and that there are no really common species. The four samples are rather

similar and there is not the expected higher diversity in the two tropical

samples.

Every ecologist knows that the diversity of species of plants and

animals is much higher in the tropics than in the temperate regions, and that

there is a latitudinal gradient of decreasing diversity from the equator

towards the polar regions. This is certainly true of trees, birds, butterflies

and a few other groups that have been well studied, but on the basis of the

figures in Table | is not true of the Ichneumonidae where all four samples

are of the typical tropical type: many species but none particularly com-

mon. What is the explanation?

First, a word of caution because it is obviously necessary to obtain

more comparable samples of Ichneumonidae to ascertain whether what has

thus far been discovered is a general phenomenon. It would also be valuable

to have temperate and tropical samples of other Hymenoptera that are para-

sitoids of insects. It might be that in the tropics other families of para-

sitoids, including dipterous families, are more diverse than in the temperate

regions.

D. Owen: A hole in a tent Sa

Another problem is seasonal heterogeneity in samples taken over an

entire year. Ecological conditions are constantly changing and there is the

possibility that the cumulative number of species is higher in a tropical

environment than in a temperate one. In view of this I have re-examined the

Kampala and Leicester samples to see how many species were added each

month during the sampling period.

The Kampala year started in July 1964 when 81 species were taken. In

August there were 58 species, 53% of which did not occur in July. Of the

47 species taken in September, 36% were new to the sample. Thereafter

the percentage of species added until April 1965 varied between 21% and 29%

per month, which suggests a relatively stable rate of addition of species.

In May 1965, 59 species were taken, 15% of them additions, and in June, the

last month of the sampling period, there were 57 species, 12% of them ad-

ditions to the total sample. As expected the rate of addition was highest

in the first few months; it then seemed to stabilize, and in the last two

months began to fall. Clearly it would have been worth continuing sampling

for a few more months.

The Leicester year started in January 1972 but no ichneumonids were

caught until the first warm weather in March when three species were taken.

In April there were six species, three of them additions. And then 62

species were taken in May, 92% of them additions which, of course, is a

simple sampling effect resulting because so few species had been taken in

the previous months. By June ichneumonids were much more abundant and

116 species were taken, 69% of them additions. In July 152 species were

taken, 51% of them additions, while in August, the best month, there were

183 species, 33% additions. Thereafter the number of species fell off with

18% added in September, 14% in October and only 4% in November when only

25 species were taken as the season was almost over. None was taken in

December.

It is not easy to compare seasonal heterogeneity at the two sites be-

cause at Kampala ichneumonids occur all year round with no obvious peaks

of abundance and diversity while at Leicester both abundance and diversity

are strongly seasonal. It is, however, obvious that at both sites the species

composition changes markedly from month to month. Ichneumonids were

tabulated from the Leicester trap for the following year, 1973, and a further

129 species were added; only 255 of the 455 species in the two years oc-

curred in both years (Owen, Townes and Townes 1981).

When the two tropical and temperate samples were first compared

(Owen and Owen 1974) it was suggested that the similarity between them is

because ichneumonids are niche specific rather than host specific. This sug-

gestion implies that there are no more niches for ichneumonids in the tropics

than in temperate regions, and is derived from a general statement in Townes

(1972b) for the family as a whole. Townes (1972b) writes, "Ovipositing adult

parasites are attracted not to hosts, but first to ecological niches and after

that to whatever hosts might be present. Host species that do not occur in

the micro-habitat of the exploring female parasite are never attacked ...

A parasite of pupae in leaf rolls will not attack pupae in cracks in the bark

or on the ground. Very seldom do parasites attacking hosts on grasses turn

to hosts on forbs, or do parasites of hosts on conifers attack hosts on

38 Contrib. Amer. Ent. Inst., vol. 20, 1983

deciduous trees. The micro-habitat of the ovipositing female can be very

specific, but within its own micro-habitat it may attack a large variety of

hosts. Thus, to say that a parasite attacks a particular host is an incomplete

statement. One should say that a parasite attacks hosts in a certain micro-

habitat, among which is the species in question." This statement is not

based on detailed observations of ichneumonids seeking micro-habitats and

hosts in the wild but from rearing records. As Townes (1972b) points out,

field observation is virtually impossible because most ichneumons are difficult

to identify to species in the field.

I am convinced that the Ichneumonidae are an established exception to

the rule of increasing species diversity from high to low latitudes, and I am

equally convinced that the rule may not be as general as has been widely

assumed. The difficulty is that most studies of tropical versus temperate

diversity have been made on relatively few groups of conspicuous and well-

known organisms like birds and trees. What is needed are comprehensive

data for other groups such as non-woody flowering plants which (excluding

epiphytes) might turn out to be just as diverse in temperate as in tropical

forest; noctuid moths might also repay study.

MONITORING BUTTERFLY ABUNDANCE

Perhaps because there are only about sixty species, much interest and

concern is directed towards Britain's butterfly fauna. Most agree that many

species have declined in recent years and blame modern farming methods and

the replacement of deciduous woodland with plantations of non-native coni-

fers. One problem is that every so often there is a good year for butter-

flies which casts doubt on whether farming and forestry have had the repeat-

edly claimed bad effects on the fauna. Another problem is that butterfly

abundance is difficult to monitor. Various techniques have been tried, includ-

ing transect counts and capture-mark-recapture, but with limited success.

The Malaise trap in the Leicester garden has been operated at the same

site for ten consecutive years and has yielded a wealth of information on

annual and seasonal variation in the numbers and diversity of many groups of

insects. As each year is completed the information obtained becomes more

and more valuable. It is easy to see that there is no such thing as a normal

year and that the populations of many of the commoner species are subject

to enormous fluctuations in numbers.

Twenty-one species of butterflies have been recorded in the Leicester

garden. Numbers vary from year to year but since the garden butterfly com-

munity is highly mobile (few species actually breed there) it is difficult to

assess whether there are good and bad years. The Malaise trap has taken

relatively few butterflies but numbers are sufficient to demonstrate annual

variations in butterfly activity, as shown in Table 2.

Thirteen species were trapped. The annual total of individuals varied

from 4 in 198] to 172 in 1973. The trap confirms that numbers vary markedly

from year to year and it is probably an effective monitor of butterfly ac-

tivity which could be useful in more natural habitats where both numbers and

the variety of species are higher.

D. Owen: A hole in a tent 39

TABLE 2. Annual variation in numbers of butterflies in a Malaise trap in a

garden at Leicester, England, 1972-1981.

1972 1973 1974 1975 1976 1977 1978 1979 1980 1981

Thymelicus

sylvestris — ~ ~ ~ 1 - - - fe e

Ochlodes venata 1 1 _ _. - - ti it = fe

Gonepteryx rhamni- - 1 1 ~ ms i he Be &. J

Pieris brassicae 2 9 12 20 2 a 1 7 4 -

P. rapae 24 115 41 58 44 5 33 13 10

P. napi 1) 123 20 14 12 6 25 6 8

Anthocharis

cardamines - _ - _ _ 1 2 1 3 x

Lycaena phlaeus ~ ~ 3 ~ _ - - Z -

Vanessa atalanta ~ 1 ~ _ 2 1 “ 6 tt

Aglais urticae 3 18 3 1 = = ee i &

Inachis io 1 Z _ 1 = ae oe re @

Lasiommata

megera 1 1 - 4 6 a ‘ee os a ts

Maniola jurtina 1 1 ~ ~ 10 i ~ - cs -

Total 43 172 77 =103 79 «14 61 LP as 4

In the tropics Malaise traps catch substantial numbers of butterflies.

The trap in the Kampala garden took 3661 in a year and the trap in the

Freetown garden took 899. The specimens were separated into families, but

in most cases not into species, but it was clear that had there been time to

do the necessary taxonomic work an impressive picture of seasonal changes

in abundance and diversity would have emerged.

TWO TRAPPING METHODS COMPARED

Malaise traps take large numbers of small moths but at most sites

rather few of the larger species are caught. On 22 June 1961 the trap

operated on Evans Old Field in Michigan caught a specimen of the medium-

sized sphingid, Xylophanes tersa. The species had not appeared. in mercury

vapor light traps operated in the vicinity and, moreover, had only once be-

fore been recorded from Michigan (Moore 1955). I put this down as an

interesting record and thought no more about it; after all to sample

Sphingidae the most effective method is to use a light trap.

40 Contrib. Amer. Ent. Inst., vol. 20, 1983

Between October 1966 and May 1968 I operated a mercury vapor light

trap in the garden at Freetown, Sierra Leone, and caught 6619 Sphingidae of

52 species. Four Malaise traps were operated continuously (day and night)

in the same area between January 1967 and May 1968 and produced 221

Rms of 29 species. Full details of these two samples are given in Owen

1969a).

The relative frequency of the species of Sphingidae in the two samples

is markedly different. All but two of the species caught in the Malaise traps

were taken in the light trap. One is a day-flyer and would therefore not be

expected in the light trap. For nearly all the commoner species the light

trap was much more effective than the Malaise traps. The Malaise traps had

therefore contributed rather little to the species list for the area.

Both samples contained a few common species and many relatively rare

species enabling oC , the logarithmic index of diversity (Fisher, Corbet and

Williams 1943) to be calculated. The value of oC =diversity for the light trap

sample is 7.9 + 0.4 and the value for the sample from the Malaise traps is

8.9 + 1.1. The two values are not significantly different. Hence despite

totally different trapping methods and different relative frequencies of

species, the values of overall diversity for the site are not different. In this

instance one trapping method would have sufficed to obtain a diversity es-

timate for the local Sphingidae.

DETECTING INSECT MOVEMENTS

Many insects periodically undertake mass movements from one place to

another, often in response to a change in weather or a failure in the local

food supply, or both. The meadow spittlebug, Philaenus spumarius , invades

Evans Old Field when surrounding alfalfa fields are mown, an action which

quickly deprives the bugs of food (Evans and Lanham 1960). Malaise traps

are highly effective monitors of insect movements, often picking up the be-

ginnings of an influx into an area before it has been noticed by even the

most observant entomologist.

The ten years of Malaise trapping in the Leicester garden have provided

numerous examples of movements into the area, particularly of Syrphidae

(Owen 1981), Coccinellidae, and certain species of moths. Nearly always the

weekly samples from the trap gave the first indication of a movement into

the garden which, for certain species in certain years, later produced immense

numbers. The Leicester trap is particularly effective at catching Coccinel-

lidae. The annual catches of three of the nine species recorded in the trap

are given in Table 3.

Adalia 2-punctata is normally the commonest coccinellid in the garden

where both larvae and adults are predators of aphids feeding chiefly on woody

shrubs and trees. As shown in Table 3, the numbers taken each year vary,

but not conspicuously, except in 1979 when the population collapsed and 1981

when again the species was not common. In this species there is no evidence

of influxes and the fluctuations in numbers are believed to be the result of

events in the garden itself. It is known, for example, that numbers of certain

species of aphids vary considerably and this alone could affect the survival

rate of both larval and adult A. 2-punctata.

D. Owen: A hole in a tent 41

TABLE 3. Annual fluctuations in numbers of three species

of Coccinellidae in a Malaise trap in a garden

at Leicester, England, 1972-1981.

—_——

Adalia Coccinella Coccinella

2-punctata 7-punctata 11~punctata

1972 168 4 2

1973 324 _ a

1974 155 ~ 1

1975 424 121 108

1976 346 564 98

1977 359 86 12

1978 La 33 es

1979 30 7 ~

1980 129 18 ~

1981 ee 2 “

For the first three years, Coccinella 7-punctata was rare (1972) or

absent (1973 and 1974) in the Malaise trap catches. This species is more

characteristic of open country where larvae and adults feed on aphids on

herbaceous plants, including field crops. In the first two weeks of August

1975 large numbers entered the garden during a period of exceptionally dry

weather. It was presumed at the time that this influx was caused by a

crash in numbers of aphids in nearby fields, perhaps brought about by the dry

weather. The influx was followed by successful overwintering in the garden

and considerable numbers in the trap in the early part of the 1976 season.

Then in the third week of July 1976 even larger numbers came into the gar-

den and stayed on to overwinter. This influx occurred when conditions were

particularly dry and again it was believed to have originated from surrounding

fields. Indeed the 1976 summer was the driest and sunniest in southern and

central England for about 250 years, a fact which was repeatedly shown in

Malaise trap records of rare and unusual insects which, presumably, were

moving about more than usual. From 1977 onwards there were no further in-

fluxes and the relatively high numbers (compared to 1972-1974) are attributable

to the species breeding in the garden. It was not until 1979 that the effects

of the 1975 and 1976 movements into the garden had disappeared.

Coccinella 11!-punctata is in Britain essentially a coastal species and

in most years is rare or absent in the Leicester Malaise trap, as shown in

Table 3. In 1975 large numbers entered the garden at the same time as the

influx of C. 7-punctata, some of them, it was believed, having travelled

42 Contrib. Amer Ent. ist., vols 20, 1983

considerable distances. They did not breed and by 1977 had become rare.

Only two were recorded in the trap in 1978, and none since.

The resident garden community of coccinellids consists essentially of

A. 2-punctata and a sprinkling of five other species, including variable num-

bers of Propylea 14-punctata which may at times invade from surrounding

woodland. The 1975 and 1976 influxes of C. 7-punctata and C. 11-punctata

disrupted the community by altering the species composition. The Malaise

trap faithfully timed and monitored these influxes in a way that would not

have been possible by eventhe most careful observations.

QUANTIFYING INSECT MIGRATION

Every year the plusiid moth, Autographa gamma, arrives in Britain from

the continent of Europe and breeds successfully. It is doubtful if it ever

successfully overwinters in the wild at any stage of its development. Al-

though direct evidence is scarce, it seems likely that substantial numbers of

A. gamma fly south from Britain in autumn. Table 4 shows the date of first

arrival of immigrant A. gamma and the number taken in the Malaise trap in

the Leicester garden during ten consecutive years. As shown, the moth nor-

mally arrives in June; its early arrival in May 1976 can be attributed to the

exceptionally dry weather in western Europe that year. Numbers fluctuate

markedly: 1973 and 1975 were "good years", and 1981 was a very poor year.

These figures confirm that numbers of A. gamma vary from year to year but

they do not provide information on spring and autumn flight directions be-

cause the trap is not designed to separate insects entering from different

directions.

TABLE 4. Date of first arrival and fluctuations in

numbers of the migratory moth, Autographa

gamma, in a garden at Leicester, England,

1971-1981.

oe ate OF. kg, Min ek taken

arrival in Malaise trap

1972 ? Ah

1973 16 June 233

1974 4 June 20

1975 30 June 155

1976 22 May 51

1977 25 June 37

1978 3 June 23

1979 10 June 17

1980 6 June 24

1981 16 June A

D. Owen: A hole in a tent 43

Malaise traps that collect and separate insects arriving from different

directions have been used to monitor Trichoptera flying up and down over a

small stream in southern Sweden (Svensson 1974). The Malaise trap is of

course an interceptor and there is no reason why suitable modified designs

should not provide information on the flight directions of insects involved in

long-distance migratory flights. Such traps have been designed and used to

quantify flight directions taken by butterflies migrating through Gainesville,

Florida (Walker 1978).

These traps are much larger than the light-weight trap designed by

Townes (1972a). They are erected in places where butterflies are known to

migrate and if run for a year provide information on numbers moving north

and south. Thus Walker (1978) demonstrated a substantial southerly movement

in authumn of the hesperiid, Urbanus proteus, and similar but less substan-

tial movements of seven other species of butterflies. With so many butter-

flies flying south in autumn a return flight north in spring is to be expected,

but except for a small movement of the nymphalid, Precis coenia, no

evidence of this was obtained from the Malaise traps. As Walker (1978)

points out, there may be several reasons why the northward flight was not

effectively monitored by the Malaise traps.

The most obvious is that the northward flight involves many fewer in-

dividuals. Observations of migratory butterflies in many parts of the north

temperate region suggest that almost invariably many more individuals are

seen flying south in autumn than north in spring (Williams 1930). Another

reason is that northward flights may tend to occur too high for the trap to

intercept them. And a third reason is that an insect entering from the north

has a greater chance of being trapped than one entering from the south.

Insects move towards the light and since at Gainesville the sun has a souther-

ly aspect, butterflies arriving from the north might have a greater chance of

ending up trapped. Walker (1978) tested this possibility by comparing numbers

of the non-migratory hesperiid, Hylephia phyleus , and found no statistical

difference between north and south samples.

It is the ability of the Malaise trap to continually monitor, by day and

night and in all weather, that makes it superior to most other trapping

methods for flying insects. For assessing the volume of butterfly migration

it is much easier to use a trap than to stand still for hours counting individ-

uals as they fly past. Not all butterflies are immediately identifiable in the

field and this is another reason why observation has its limitations.

UNDERSTANDING SEASONAL CYCLES IN EQUATORIAL VERTEBRATES

Kampala at 0° 20' N is an ideal place for the study of the seasonal

breeding and migratory cycles of mammals and birds. The temperature changes

little during the year, fluctuations in each 24-hour period being greater than

seasonal changes, and daylength is effectively constant. Rainfall occurs in all

months of the year with two seasonal peaks which, however, are not reliable.

It is an extremely uniform climate but despite this most vertebrates that have

been studied exhibit seasonality in their breeding or migratory cycles. Thus

Mutere (1973) found seasonal peaks in the breeding of two species of insec-

tivorous free-tailed bats, Tadarida, which are correlated to some extent with

44 Contrib. Amer. Ent. Inst., vol. 20, 1983

seasonal changes in the amount of rainfall. He postulated that rainfall acted

as a proximate stimulus for breeding and that variation in the availability of

night-flying insects, particularly moths, is the chief ultimate factor. But al-

athough rainfall figures were easy to obtain, information on seasonal fluc-

tuations in abundance of night-flying insects (the food of the bats) was un-

available, with the notable exception of the results from the Malaise trap in

the Kampala garden. The trap indicated that during a year the mean number

of moths caught per 24 hours varied between 196 (May) and 73 (August).

This provided the only information on possible variation in moth abundance

which could be correlated with bat breeding cycles.

Many insect-eating birds that breed in the Palearctic fly south in

autumn and "winter" on or near the equator in Africa. There is much cir-

cumstantial and some experimental evidence that the stimulus to commence

the southward migration is provided by decreasing daylength, decreasing

temperature, and a progressive failure of the insect food supply. But how do

these birds time the commencement of their return flight in February and

March? On the equator daylength and temperature are effectively constant

and could hardly act as stimuli for the birds to set off on the long north-

ward flight back to the breeding areas.

Because of the apparent lack of a seasonal trigger, it was postulated

for one species, the yellow wagtail (Motacilla flava), which in the northern

winter is extremely abundant around Kampala, that there is an internal timer

which (soemhow) is set to act at the correct time for the northward de-

parture (Curry-Lindahl 1958, Marshall and Williams 1959). Evidence for the

existence of an internal timer is based on a failure to find environmental

stimuli that might trigger the start of the northward flight. Such evidence

is to say the least rather unsatisfactory. The possibility that insect numbers

on or near the equator fluctuate sufficiently on a seasonal basis to stimulate

the return migration of the yellow wagtail was discounted (Marshall and

Williams 1959) or ignored (Curry-Lindahl 1958).

Table 5 shows the mean number of insects per 48 hours in the Kampala

Malaise trap for each month of the year starting July 1964. It also shows

the actual rainfall for each month and the 10-year mean monthly rainfall.

Monthly variations in insect numbers correlate better with the mean "expected"

rainfall than with the actual rainfall, suggesting that in many species there is

a built in cycle with the peak abundance at the best time of year, during and

just after the annual rainfall peak. Yellow wagtails begin to leave Kampala

in February, the least good month for insects, and continue to leave through

March and early April, when insect numbers are increasing towards the May

peak. Their departure thus occurs during the period of maximum change in

the abundance of their potential food supply, and this itself could provide

the necessary stimulus to migrate north.

The figures. in Table 5 are based on only one year of Malaise trap sam-

pling. Caution is therefore necessary in interpreting fluctuations in numbers

as being sufficient to provide the stimulus for yellow wagtails (and other

insectivorous birds) to set off for the north, but the possibility remains, es-

pecially as essentially the same evidence of fluctuations in abundance was

obtained by sweep sampling on the Athi Plains near Nairobi, Kenya (Dingle

and Khamala 1972).

D. Owen: A hole in a tent 45

TABLE 5. Seasonal fluctuations in insect numbers and rainfall,

Kampala, Uganda, July 1964-June 1965. From Owen

(1969b).

ta wn, ‘thy: Mean mumbet. oc detuel 9. lO -yeer mean.

of insects per rainfall rainfall, 1953-

48-hour sample (mm.) 1962 (mm.)

July 669 24 51

August 480 85 111

September 452 206 98

October 399 135 121

November 436 140 115

December at 216 96

January 458 15 64

February 196 73 53

March 668 139 139

April 1156 159 174

May 1176 50 95

June 790 10 61

I was not of course able to show that fluctuations in insect numbers

directly affect the availability of food to the birds, but the existence of such

fluctuations must be taken into account before the concept of the internal

timer is accepted. My hypothesis makes fewer assumptions in the field of

avian physiology than that of Curry-Lindahl (1958) and Marshall and Williams

(1959) and for this reason alone it is to be preferred until more evidence is

forthcoming.

THE HOLE IN THE TENT

I have been Malaise trapping for more than twenty years and | still find

it absolutely compulsive, whether in some remote tropical place or in my own

garden. Indeed the garden trapping, now in its eleventh year, has produced

a quite extraordinary amount of information on the abundance and diversity

of many groups of insects. It has also been exciting, even though nothing

quite as unusual as Owenus has turned up, and the trap continues to produce

new records of insects for this part of England.

I must acknowledge Rene Malaise's perception when he realized what

was going on at the hole in the top in his tent. I acknowledge even more

the efforts of Henry and Marjorie Townes in developing Malaise traps and for

the enormous amount of taxonomic work they have put into my samples of

Ichneumonidae and other parasitic Hymenoptera. It is an honor to dedicate

this paper to Henry Townes on the occasion of his 70th birthday.

46 Contrib. Amer. Ent. Inst., vol. 20, 1983

REFERENCES

Curry-Lindahl, K. 1958. Internal timer and spring migration in an equatorial

migrant, the Yellow Wagtail (Motacilla flava). Arkiv. Zool. 11:

541-557. |

Dingle, H. and Khamala, C.P.M. 1972. Seasonal changes in insect abundance

and biomass in an East African grassland with reference to breeding and

migration in birds. Ardea 59: 216-221.

Evans, F. C. and Lanham, U. N. 1960. Distortion of the pyramid of numbers

in a grassland insect community. Science 131: 1531-1532.

Evans, F. C. and Murdoch, W. W. 1968. Taxonomic composition, trophic

structure and seasonal occurrence in a grassland insect community.

J. Anim. Ecol: 37: 259-273.

Evans, F. C. and Owen, D. F. 1965. Measuring insect flight activity with

a Malaise trap. Pap. Mich. Acad. Sci. Arts Letters 50: 89-94.

Fisher, R. A., Corbet, A. C.’and Williams, C.B. 1943. The relation between

the number of species and the number of individuals in a random sample

of an animal population. J. Anim. Ecol. 12: 42-58.

Malaise, R. 1937. A new insect-trap. Ent. Tidskr. 58: 148-160.

Marshall, A. J. and Williams, M. C. 1959. The pre-nuptial migration of the

Yellow Wagtail (Motacilla flava) from latitude 0.04' N. Proc. Zool.

Soc. Lond. 132: 313-320.

Moore, S. 1955. An annotated list of the moths of Michigan exclusive of

Tineoidea (Lepidoptera). Misc. Pub. Mus. Zool. Univ. Mich. 88: 1-87.

Mutere, F. A. 1973. Reproduction in two species of equatorial free-tailed

bats (Molossidae). E. Afr. Wildl. J. 11: 271-280.

Owen, D. F. 1969a. Species diversity and seasonal abundance in tropical

Sphingidae (Lepidoptera). Proc. Roy. Ent. Soc. Lond. A. 44: 162-168.

Owen, D. F. 1969b. The migration of the yellow wagtail from the equator.

Ardea 57: 77-85.

Owen, D. F. 1971. Estimates of species diversity in tropical Ichneumonidae.

Rev. Zool. Bot. Afr. 83: 173-177.

Owen, D. Fo and Chanter;°D.°O." 1970. “Species diversity and seasonal abun-

dance in tropical Ichneumonidae. Oikos 21: 142-144.

Owen, D. F. and Owen, J. 1974. Species diversity in temperate and tropical

Ichneumonidae. Nature 249: 583-584.

Owen, J. 1981. Trophic variety and abundance of hoverflies (Diptera,

Syrphidae) in an English suburban garden. Holarctic Ecol. 4: 221-228.

Owen, J., Townes, H. and Townes, M. 1981. Species diversity of

Ichneumonidae and Serphidae (Hymenoptera) in an English suburban

garden. Biol. J. Linn. Soc. 16: 315-336.

Steyskal, G. C. 1981. A bibliography of the Malaise trap. Proc. Ent. Soc.

Wash. 83: 225-229.

D. Owen: A hole in a tent 47

Svensson, B. W. 1974. Population movements of adult Trichoptera at a

south Swedish stream. Oikos 25: 157-175.

Townes, H. 1962. Design for a Malaise trap. Proc. Ent. Soc. Wash. 64:

253-262.

Townes, H. 1970. The genera of Ichneumonidae. Part 2. Mem. Amer.

Ent. Inst. 12: 1-537.

Townes, H. 1972a. A light-weight Malaise trap. Ent. News 83: 239-247.

Townes, H. 1972b. Ichneumonidae as biological control agents. Proc. Tall

Timbers Conf. Ecol. Anim. Contr. Hab. Man. 3: 235-248.

Walker, T. J. 1978. Migration and re-migration of butterflies through north

peninsular Florida: quantification with Malaise traps. J. Lepid. Soc.

32: 178-190.

Williams, C. B. 1930. The migration of butterflies. Oliver and Boyd,

Edinburgh.

A NEW NEARCTIC SPECIES OF HADRODACTYLUS

(HYMENOPTERA: ICHNEUMONIDAEBE)

Mattias Idar

Department of Entomology, University of Uppsala

S-751 22 Uppsala, Sweden

Hadrodactylus townesi, n. sp.

Hadrodactylus townesi_ is very similar to the palearctic H. semirufus

(Holmgren) (Idar 1979, Fig. 8 A-L, 1981:236-237). It differs essentially by

more shagreened metapleurum and by extensive reddish coloration on head

and thorax. [Ocelli and hind coxae are not diagnostic as it is essentially as

in faciator (Thunberg) and vulneratus (Zetterstedt) in both the above

species ]. Such extensive reddish coloration is not observed in any Palearctic

forms of the genus. It can be recognized from other Nearctic species with

reddish thorax by its larger ocelli, more slender petiolar segment, hind coxa,

and hind femur.

Head rather strongly narrowed backwards; lateral ocellus diameter equal

or subequal to the distance to the other; cheek length about 0.3 the width

of mandible basally; mandibular teeth essentially equal in length and width;

epomia indistinct in 80% of the specimens; meropleurum with very indistinct

or no wrinkles in upper anterior portion; metapleurum shagreened between

the punctures; propodeum with apical area closed anteriorly and medially;

petiolar segments slender, maximum width about 0.25 of length, with weak

median dorsal ridges in 25% of the specimens; hind coxamore than twice

as long as wide; hind femur more than five times as long as wide; fifth

hind tarsal segment about 1.5 as long as the fourth.

Body color yellowish-red; area between ocelli and antennae, edges of

propleurum, frontal edges of pronotum, and anterior surface of mesosternum

dark to blackish-brown; propodeum laterally, petiole, and tergites II-VI more

or less marked with reddish-brown; face, inner orbit, clypeus, mandible,

palpi, antennal base ventrally, hind corner of pronotum, tegula, and fore and

middle trochanters yellow or yellowish. Males also with marking on meso-

sternum (as in faciator) and fore and middle coxae yellow.

Holotype: Female, Mt. Madison, N.H., June 26, 1938, H. & M. Townes

(CPH,Gainesville, FL.). Paratypes: 1 M, same data as holotype; 2 F,

Stoneham, P.Q., June 18, 1938, H. & M. Townes; | F, Pinkham Notch, N.H.,

June 25, 1938, H & M. Townes; IF, Huron Mts., Mich., June 21, 1961, H. &M.

Townes. All paratypes are in the Townes Collection except for one F from

Stoneham in my collection.

REFERENCES

Idar, M. 1979. Revision of the European species of the genus Hadrodactylus

Forster (Hymenoptera: Ichneumonidae). Part 1. Ent. Scand. 10: 303-313.

idat, My.196L.. Same. Part 2. Ent, Scand, 12: 231-239,

A NEW SOUTH AFRICAN SUBFAMILY RELATED TO

CARDIOCHILINAE (HYMENOPTERA : BRACONIDAE)

W.R.M. Mason

Biosystematics Research Institute

Agriculture Canada, Ottawa, KIA 0C6

Abstract

Two new genera and 6 new species are described and placed in a new

subfamily of Braconidae (Hymenoptera). A cladistic scheme is proposed re-

lating the new forms to Cardiochilinae, Microgastrinae and Miracinae.

Recently I found specimens of an undescribed subfamily related to

Cardiochilinae among unnamed South African material in the Townes Collec-

tion. It is a pleasure to dedicate this study to the great collector, cataloguer

and taxonomist.

SUBFAMILY KHOIKHOIINAE, new subfamily

Thick-set species resembling Cardiochiles but with a longer abdomen.

Color mostly blackish with some reddish areas and brown wings.

Head weakly transverse, temples in dorsal aspect about as long and wide

as eyes; frons concave; scape truncate and not tapered apically; flagellum of

medium length, between 20 and 40 articles, the articles cylindrical and

tightly fitted except for the antennal apex of females where articles are sub-

moniliform; clypeus small, its apical margin weakly concave or prolonged in-

to a large conical point; labrum semicircular, hairy and conspicuous; mandi-

ble teeth twisted about 45°, the anterior tooth much the larger; occipital

carina absent; maxillary palpi 5-jointed, the basal 2 fused; labial palpi 4-

jointed.

Median part of pronotum flat, lorate, and with no special median mod-

ifications; lower outer corner of propleuron not overlapping the pronotum;

lateral triangles of pronotum much longer than tall; notauli deep and

crenulate, meeting far in front of scutellar scrobe; a shallow parapsidal fur-

row extending forward from axilla region; transcutal suture strong and com-

plete, weakly concave anteriorly and running clearly in front of the

scutellar scrobe; behind transcutal suture two flat, well defined axillae half

or more as large as scutellum; polished apical margin of scutellum inter-

rupted medially by a rugulose area; prepectal carina absent; propodeum

generally rugose with a more or less well-defined medial longitudinal carina;

tarsal claws simple, hind tarsi with no median ventral ridge of appressed

hairs but the hind basitarsus large and compressed.

Wing structure (Fig. 16) resembling that of Cardiochiles; vein A2 of

forewing strong and reaching the margin; vein A3 strongly pigmented, though

without defined anterior edge, running along posterior margin to meet A2

_and form an enclosed anal cell; pigmented but indefinite transverse vein (r)

in radial cell of hind wing closer to apex than to base of that cell; veins 1A

and cu-a of hind wing meeting at an obtuse angle.

First tergite of metasoma clearly delimited on both sides, usually barrel-

shaped, the length and maximum width about equal; surface of tergite

roughened and bearing a deep median groove; spiracle of first segment in a

50 Contrib. Amer. Ent. Inst., vol. 20, 1983

large lateral membrane that is evenly covered with fine parallel striations and

contains no laterotergite; sternite 1 rectangular, not divided into the anterior

and posterior plates usual in Ichneumonoidea but appearing more heavily

sclerotized anteriorly. Tergites 2 and 3 rectangular and about equal, fused

but mutually delimited by a straight suture; remaining abdominal segments

all visible; cerci appearing as oval plates on hind margin of metasomal tergum

8; metasomal sternum 9 clearly visible apically in both sexes; sternite 7 of

males deeply emarginate apically; hypopygium of female entire, medially ob-

tusely or acutely pointed; ovipositor short and scarcely visible or about as

long as abdomen.

Members of the Khoikhoiinae bear a general likeness to Cardiochilinae

and will run there in present braconid keys, because of the anteriorly convex

arch of the 3Rs vein, wider than high 1Rs cell, many-jointed antennae,

"sessile abdomen, etc. They have practically the same venation and anten-

nal structure as Cardiochiles but the following couplet will separate them.

Metasomal tergite 1 bearing a Y-shaped groove with a rounded elevated area

apically between the arms of the Y (Figs. 20, 21); laterotergite 1

present but rarely differentiated distally from median tergite; transcutal

groove absent; clypeal margin convex and often bearing 2 small toothlets,

labrum almost completely concealed (World wide) . . . CARDIOCHILINAE

Metasomal tergite 1 with only one deep, median groove (Figs. 4, 15); latero-

tergite 1 absent, the pleural areas soft and closely striate with the

spiracle in the membrane (Fig. 9); transcutal groove strong, lying in

front of prescutellar scrobe (Fig. 3); clypeal margin concave and some-

times bearing a large median tooth, labrum conspicuously exposed (South

PCA) Neste Coe ra rine a at pion K HOIK HOIINAE

KEY TO SPECIES OF KHOIKHOIINAE

1. Clypeal margin concave (Fig. 5); face smooth, at most punctate, flagel-

lum with less than 30 articles; frons centrally smooth; female hypo-

pygium extending far beyond tergum 8; ovipositor sheath thin and

about as long as abdomen (Pied) #-8)).0 04 Sania, pp wen.. : 2

Clypeal margin with a large medial point (Fig. 13); face dull and coarse-

ly rugose; flagellum with over 30 articles; frons completely trans~

versely striate and medially carinate; female hypopygium ending about

at abdominal apex; ovipositor scarcely protruding (Figs. 9-15) .....

Aug ie 2 eee Ws Da ee eg ee KRhOikHoia, oN. Sen... « -/<:4

2. . Mesopleutal suture about 3/4 as dong as pleuwron i)... ob ck 6 2 3

Mesopleural suture only about 1/4 as long as pleuron............

3. Mesoscutum strongly punctate; upper half of metapleuron smooth; ab-

dominal terga 2-5 mostly smooth and shining. . 2. S. henryi, n. sp.

Mesonotum smooth; metapleuron strongly rugulose; abdominal terga 2-5

mostly dull, coriaceous-rugulose ....... Lo Si omaryoriae; i on. sp.

4. Mesonotum and mesopleuron densely and coarsely punctate ........

Se a OSS SS a oe Sieh ee ea BPO cee 1. Ku Cownesi, n. sp.

Mason: Khoikholinae om

Mesonotum and mesopleuron smooth, finely or sparsely punctate .... 5

5. Wings almost uniform brown from base to apex; flagellum with 32-33

Brvicies: Verie™ SINGo oe ee ea 2 i solata, i. Sp.

Wings hyaline basally, only brown beyond the stigma; flagellum with

about 37 articles; vertex coarsely punctate . 3. K. semiadusta, n. sp.

SANIA, ‘new genus

(Figs. 1-8)

Type-species: Sania marjoriae Mason

Differs from the subfamily description as follows: flagellum with 24-

28 articles; frons behind antennae excavated and medially mostly smooth,

transversely striate only at edges; clypeus protuberant, over half as wide as

face, its apical margin concave; clypeus and face smooth and bearing a broad

longitudinal, median swelling; temple (in dorsal aspect) slightly longer than

eye, convex, but only as wide as, or narrower than, the eyes; lateral carina

and groove of axilla usually weak or absent; metasomal sternites 2-4 of fe-

male wide and rectangular, sternite 5 of female strongly emarginate, only

visible as a pair of lateral triangles; female hypopygium acutely pointed, much

longer than deep (in lateral aspect) and extending far beyond the dorsal ab-

dominal apex; sternite 7 of male deeply emarginate; ovipositor sheaths very

think, extending far beyond the abdominal apex and curved upward, hairs

sparse except for an apical clump; length about 4 mm, fore wing 3 mm.

1. Sania marjoriae, n. a

Holotype, female: Flagellomeres 28, the basal ones twice as wide as the

apical, the apical 8-10 articles submoniliform; antenna about 0.9 as long as

fore wing; inner and ventral apical margins of scape with a concave marginal

shelf, behind which apical margin of scape is weakly flared out; clypeus

smooth and finely punctate; fact mostly weakly transversely aciculorugulose;

frontal calli rugulose; vertex smooth; temples punctate and roughened.

Side of pronotum completely rugose; lobes of scutum, scutellum and axil-

lae convex, smooth and very finely punctate; hairs of central lobe of scutum

all pointing backward, not angling outwardly; mesopleuron smooth, furrow

long and deep but ending well before the mid coxa; propodeum with several

irregular transverse subbasal rugae.

Tergite 1 of metasoma about 10% longer than wide, the widest point

near the apical 0.2, thus mostly tapered toward base; median groove deep but

very wide; surface of tergite 1 mostly rugose but irregularly coarsely acicu-

late medially; remaining terga, and also the sterna, dull and weakly,

tA feminine generic name, dedicated to the San people, a few of whom still

follow their ancient hunting life in remote and arid parts of southern Africa.

2 ‘ Resi :

Dedicated to Marjorie Townes, whose quiet labors over many years have

contributed so greatly to the production of the "Townes and Townes"

team.

52 Contrib... Amers-EBnt.cinst.y vol: 20, 1983

irregularly, regulose; groove between metasomal terga 2 and 3 very broadly .

V-shaped, concave behind; medial band on tergite 2 longitudinally finely

aciculate.

Color of body black with castaneous to fulvous suffusions on the follow-

ing parts; mandibles, cheeks, corners and collar or pronotum, mesopleuron

below furrow, subtegular ridge, metapleuron, trochanters and basal parts of

tibiae; wing membrane lightly embrowned, venation dark brown; margins of

abdominal plates usually narrowly yellowish.

Holotype: Female, South Africa, Jonkershoek, near Stellenbosch, 18

December 1970, Malaise trap operated by V. Whitehead (CPH, Gainesville,

2. Sania henryi, n. sp.

Holotype, male: Flagellomeres 24, basal ones 1.5 as wide as apical ones;

antenna about 0.8 as long as forewing; clypeus and median elevation of face

smooth and finely punctate; sides of face shallowly rugulopunctate; frontal

calli raised and coarsely rugulopunctate; vertex mostly smooth; postocciput

and temples densely and coarsely punctate.

Submarginal grooves of pronotal sidepiece irregularly costate-punctate, the

central area between them indefin itely defined and not raised; mesoscutum,

scutellum and axillae moderately punctate, the punctures separated by 1-2 x

their own diameters; hairs of median lobe near posterior half of notauli

directed laterad at angles up to 90° or even pointing a little anteriorly of

that; mesopleuron mostly smooth behind but moderately punctate in front and

below the groove; longitudinal groove transcostate, large, long and deep, reach-

ing to the mid-coxa; propodeum without a transverse carina.

Tergite 1 of metasoma about 10% longer than wide, the greatest width on

the apical 0.5-0.7, a little more tapered anteriorly, base about half as wide

as middle but apex about 2/3 as wide; surface of tergite 1 verrucose, merging

to apical aciculations; other tergites with dull smooth surfaces; digitus

cultriform, about 0.4 as wide as paramere and bearing 3-4 small toothlets on

the outer curve (Fig. 8).

Color of body black with fulvous to castaneous suffusions on the following

areas; mandibles, sides of face, frontal calli, temples and cheeks, most of

pronotum, sides of mesonotal lobes, scutellum, axillae, mesopleuron below

sulcus, subtegular ridge, metapleuron, hind coxa, trochanters, base of tibiae.

Margins of abdominal segments extensively yellowish, especially ventrally.

Wing membrane brown, veins dark brown to black.

Holotype: Male, South Africa, Garies, Cape Province, 23 Sept. 1970,

H. and M. Townes (CPH, Gainesville, FL.).

3. Sania capensis, n. sp.

Holotype, male: Flagellomeres 26, basal ones twice as wide as the apical

ones; antenna about 0.9 as long as fore wing; face and clypeus smooth and

finely but closely punctate; vertex smooth; frontal calli weakly raised and

rugulose; temple vertically aciculo-rugulose.

Side of pronotum with upper, lower and posterior submarginal transcostate

grooves, medial elevation between them smooth and polished; lobes of meso-

scutum convex, smooth and finely punctate; axilla and scutellum similar in

texture; hairs near notauli on median lobe of scutum directed outwardly at

Mason: Khoikhoiinae fs

angles up to 45°; mesopleuron mostly smooth, furrow very short, medial and

with only 2-4 transverse costulae; propodeum with a subbasal transverse

carina.

Tergite 1 of metasoma about as wide as long, its sides strongly curved,

medially about twice as wide as base or apex; all terga mostly smooth but

not strongly shining because of microsculpture; tergite 1 with weak apical

sculpture, terga 2 and 3 with areas of weak transversely trending wrinkles;

digitus curved fusiform, about 1/4 as wide as paramere (Fig. 7).

Color of body black with irregular casteneous to fulvous, suffusions on the

following areas: mandibles, pronotal collar, pronotum posteriorly, notaull,

scutellum, mesopleuron, femora and tibia; wings hyaline on basal 0.6, brown

apically; veins dark brown to black but the desclerotized apical portions light

brown; margins of most abdominal plates narrowly yellowish.

Paratypes: flagellomeres 25-27; castaneous suffusions on body varying

considerably in extent and hue.

Holotype: Male, South Africa, Jonkershoek near Stellenbosch, 18 Dec.

1970, Malaise trap operated by V. Whitehead. Paratypes : 2 Males, same

data but one of them captured 1] Jan. 1972. (CPH, Gainesville, FL.).

KHOIKHOIA," new genus

(Figs. 9-15)

Type-species: Khoikhoia townesi Mason.

Differs from the subfamily description as follows: frons strongly pro-

tuberent beside upper part of eye; frons behind antennae strongly excavated,

and bearing transverse striae and a median carina; scape with marginal flat

area apically; flagellum with 32-37 articles; clypeus small, less than half as

wide as face and protruding downward as a tapering cone concealing median

and basal parts of labrum; temple (in dorsal aspect) longer than eye and

bulging outward so that head is wider behind eye; clypeus, face and sides of

frons coarsely rugose; axilla with a strong lateral marginal carina and crenu-

late groove; metasomal sternites 2-5 of female entire, rectangular and of

nearly uniform size, hypopygium of female triangular in lateral view and not

extending beyond abdominal apex; ovipositor sheath, short thick, hairy apically

and scarcely extending beyond hypopygium; length 6-7mm, fore wing 5.5 mm.

1. Khoikhoia townesi, n. sp.

Holotype, female: Flagellomeres 35, basal: middle: apical articles::

1.5:2:1; antenna about 0.8 as long as fore wing; vertex, post occiput and upper

temples coarsely and densely punctate to rugopunctate; lower temples and

cheeks mostly coarsely vertically aciculate.

Sides of pronotum entirely coarsely rugose except for upper and lower

transcostate grooves; propleuron polished and strongly punctate; mesoscutum,

The genus name is feminine, dedicated to the original inhabitants of the

South African cape region, the Khoikhoi, who were known to the Dutch

settlers as hottentots because of their unique clicking speech.

54 Contrib. Amer. Ent. Inst., vol. 20, 1983

scutellum and axillae densely, coarsely punctate to rugopunctate; meso-

pleuron densely, coarsely punctate above, rugopunctate below.

Tergite 1 of metasoma about 10% longer than wide, the lateral margins

curving, base about half as wide as middle but apex only 10% narrower; sur-

face of tergite 1 strongly longitudinally aciculorugulose centrally, polished

basally and apically, the median groove very sharp and deep; remaining terga

dull and hairy, the groove between terga 2 and 3 straight and fine; hypopygium

obtuse, extreme apex emarginate and desclerotized; apical margins of meta-

somal sternites 2-5 straight, the sternites subequal in length.

Color of head and thorax ferrugineous with black suffusions on the follow-

ing areas; clypeus, lower medial part of face, frontal excavation, vertex, pro-

notal collar, propleura, sides of scutellum, areas around wing bases but not

tegula, metepisternum, apical part of metanotum, dorsal part of propodeum.

Cheeks and clypeus yellow around base of mandible. Abdomen very dark brown

with apical margins of terga and sterna 3-6 yellowish and membranous areas

also yellowish. Legs black with most of each tibia and upper part of coxae

ferrugineous. Scape (except for thick apical margin) and pedicel reddish,

flagellum dark brown. Wings subhyaline basally but brown distad from basal

vein; venation dark brown. |

Holotype: Female, South Africa, Grahamstown, Feb. 1972, Malaise trap

operated by Fred Gess. (CPH, Gainesville, FL.).

2. Khoikhoia Soha n. sp.

Holotype, Male: Flagellomeres 32-33, basal articles about twice as wide

as apical; antenna about as long as fore wing; vertex smooth, temples weakly

rugopunctate, the sculpture toward the cheeks coarsely rugulose.

Lower margin of side of pronotum longitudinally aciculate, central part of

pronotal side elevated and punctate; propleuron polished and finely punctate;

upper, lower and posterior grooves strongly transcostate to rugulose; meso-

scutum, axillae and front of scutellar disc smooth and finely sparsely punc-

tate; sides and apex of scutellum densely and more coarsely punctate;

mesopleuron generally smooth with fine, sparse punctures, but strongly punc-

tate near the metapleuron.

Metasomal tergite 1 about 10% longer than wide, base about half as wide

as middle and apex, not narrowed posteriorly; surface, including even the

median groove, smooth and finely punctate; remaining terga smooth and

finely closely punctate; groove between terga 2 and 3 slightly convex an-

teriorly; metasomal sternites 3-6 with straight apical margins but the 7th

sternite with a V-shaped emargination, whose arms diverge at about 90° to

one another, and which divides the sternite into 2 visible sections (Fig. 11);

hypopygium strongly emarginate; digitus evenly curved and tapering apically.

Color black with reddish to castaneous suffusions on the following parts:

mandible, cheek and temple, elevated disc and upper corner of pronotum,

sides of mesoscutal lobes and parts near notauli, axillae and sides of

di ‘

From Latin, meaning sunburned, an allusion to the color and to the fully

infuscated wings.

Mason: Khoikhoiinae 55

scutellum, mesopleural ridge, central part of mesopleuron, hind femur and

basal part of hind tibia, parts of middle femur and tibia. Area around base

of mandible and frontal calli yellowish. Folded under parts of apical mar-

gins of posterior tergites yellowish. Wing membrane brown, venation almost

black.

Holotype: Male, South Africa, Cape Province, Jonkershoek near

Stellenbosch, 18 Dec. 1970, Malaise trap operated by V. Whitehead (CPH,

Gainesville, FL.).

3. Khoikhoia semiadusta‘, n. sp.

Holotype, Male: Flagellomeres 37, basal articles about 2-1/2 times wider

than apical ones; antenna about 10% longer than fore wing; vertex and tem-

ples coarsely but not densely punctate, the punctures usually separated by

half their diameter; cheeks coarsely vertically aciculate.

Sides of pronotum centrally rugopunctate and elevated, the submarginal

grooves transcostate but the vertical groove rugulose; lower margin of pro-

notum polished and punctate; mesoscutum and axillae moderately punctate;

mesopleuron polished and irregularly, but mostly finely, punctate and bearing

a series of shallow marks indicating a longitudinal furrow.

Metasomal tergite 1 about 10% longer than wide, narrowed basally to

about half the width at center or apex; surface of tergite 1 polished, basally

finely punctate, apically rugulo-aciculate; remaining terga dully polished and

hairy with some transverse wrinkling laterally on terga 3-6; sternites 2-6

similar in size and sculpture; sternite 7 apically with a broadly V-shaped

emargination whose arms diverge at 135° from one another (Fig. 12); sternite

7 about twice as long laterally as medially; hypopygium weakly emarginate;

digitus evenly curved and tapering apically.

Color black and reddish to castaneous suffusions on the following parts:

mandible, frontal calli, upper temple, tegula, apex of middle femur, base of

hind tibia, tibial spurs. Lateroapical margins of apical terga and large patches

around mandible base yellowish. Wing membranes hyaline basad of hamuli and

stigma, apically infuscated; venation dark brown.

Holotype, Male, South Africa, Cape Province, Jonkershoek near Stellenbosch,

24 Dec. 1970, Malaise trap operated by V. Whitehead (CPH, Gainesville, FL.).

CHARACTER ANALYSIS

I have found 32 characters that seem to show phylogenetically significant

variation. Nine of them are group synapomorphs for the "microgastri" ¢,

leaving 23 characters useful for internal analysis, of which 10 are autapomorphs,

leaving only 13 that show useable synapomorphies. Five of these are triple

synapomorphs of Khoikhoiinae, Microgastrinae and Miracinae: the other 8 are

1 : : :

From Latin meaning half tanned by the sun in reference to the wings, which

are brown apically and hyaline basally.

2 e e e

I have coined a vernacular term "microgastri" to refer to the four sub-

families included in this survey because it is convenient to avoid long phrases.

56 Contrib. Amero-ent.: sts, wole7gO, 1983

distributed in 5 different combinations. I have also given reduced weight to

reductional features, for example loss of cross veins, palpal articles, etc.,

which could easily have occurred several times and thus carry little informa-

tion value. Finally this leaves 5 synapomorphs that seem strong. Two of

them, 7, straight or concave clypeal margin and 21, tergite 1 with completely

delimited sides, define the triple group mentioned above: two others, 1,

constant number of flagellomeres and 17, ventral ridge on hind basitarsus

(Mason 1981], Fig. 14), define the group of Microgastrinae plus Miracinae.

Finally one, 12, propodeum with median carina, occurs in Khoikhoiinae and

Miracinae but is of doubtful value as a synapomorph because of many occur-

rences in specialized groups of Microgastrinae and Cardiochilinae. Thus my

cladogram has the first division between Cardiochilinae and the rest, the

second between Khoikhoiinae and the rest and the third between Micro-

gastrinae and Miracinae.

Synapomorphies defining the group of Cardiochilinae + Khoikhoiinae +

Microgastrinae + Miracinae (= microgastri) plesiomorphic state in brackets:

Occipital carina absent (present); tergum 1 of metasoma with a Y-shaped

dorsal groove and a mesal tumid area between the arms of the Y (tergum

simple with no such groove); spiracle of metasoma 1 located on the folded-

under side part of tergum (spiracle located dorsally or on the fold); lateral

membranes of basal abdominal segments covered with closely parallel stria-

tions (membranes smooth); metasomal sternite 1 with basal and distal sections

fused to form a single rectangular plate (basal and distal sections distinct);

spiracle of metasoma 7 absent (spiracle M7 present): cerci flat and button-

like, on sides of tergum 8; 2nd Cu of hindwing absent (2 Cu (HW.) present):

3RS of tore wing convex anteriorly (straight or concave anteriorly).

Apomorphies defining Cardiochilinae: 9, transcutal groove reduced and

scarcely discernible (transcutal groove strong); 10, axilla reduced to a vertical

area or a carinate lobe (axilla large and horizontal); 20, a medial tergal

plate [= tergite] of metasoma | partly defined, only the basal half being

clearly separated from the weakly sclerotized lateral margin (no separately

defined plates on tergum 1); 15, r-m“ (HW.) present).

Synapomorphies defining the group of Khoikhoiinae + Microgastrinae +

Miracinae: 7, apical margin of clypeus straight, exposing a large semi-

circular labrum in the space between clypeus and mandibles (clypeal margin

convex, almost concealing labrum and mandibles); 4, maxillary palpi reduced

to 5 articles by fusion of 1 and 2 (maxillary palpi with 6 articles); 21, tergite

1 of metasoma much more heavily sclerotized than remainder of tergum, its

lateral margins fully and clearly delimited (medial and lateral parts of tergum

| similarly sclerotized and neither clearly nor completely delimited); 13 veins

lr and 3r of fore wing lost (1r and 3r present, at least as traces); 16, hamuli

reduced to 3 (hamuli 4-7).

Apomorphies defining Khoikhoiinae: 8, prepectal carina absent (present);

7, apical margin of clypeus concave or pointed (margin straight); 12, pro-

podeum with a median carina (propodeum areolate); 19, tergite | of metasoma

with a median groove (with a Y-shaped groove); 23, lateral areas of tergite

1 without laterotergite but occupies by striate membrane containing the

spiracle (lateral parts weakly sclerotized and not striate); 18, tarsal claws

simple (toothed); 15, vein r-m2 of hind wing lost (r-m2 present).

Mason: Khoikhoiinae ows

Synapomorphies defining the group Microgastrinae + Miracinae: 1, number

of flagellomeres constant (variable intra-, and inter specifically); 9, transcutal

groove absent (present); 17, hind basitarsus with a median ventral palisade of

closely appressed or fused hairs (without such a ridge).

Apomorphies defining Microgastrinae: 2, flagellum with 16 articles (with

a larger number); 3, the central flagellomeres with longitudinal placodea ar-

ranged in 2 ranks, so that the break between the ranks gives the appearance

of a false division of each flagellomere into 2 subsections (placodea arranged

irregularly; 7, clypeal margin obviously concave (margin straight); 22, basal

part of metasomal tergite | subtending a lightly sclerotized setiferous latero-

tergite that bears a spiracle (the laterotergite 1 absent or not delimited); 10,

axilla of the mesoscutum poorly defined, mostly vertical (axillae large and

horizontal).

Apomorphies defining Miracinae: 2, flagellomeres 12 (more numerous); 5,

maxillary palpi with 4 articles because of fusion of 2 of the distal 3 (with

5 articles); 6, labial palpi with 3 articles (4 articles); ll, median apical

declivity of the scutellum with a pair of side by side foveae (without such

foveae); 12, propodeum with a median carina (propodeum areolate); 23,

laterotergite of metasoma | absent, spiracle lying in a striate membrane

(spiracle in sclerotized lateral part of tergum 1); 18, tarsal claws simple

(claws pectinate); 14, 15, veins rl and r-m2 of hind wing absent (these veins

developed).

NOTES ON SOME APOMORPHIES

Most of the apomorphies listed above are simple enough in structure and

polarity to need no explanation but a few call for more comment.

8, Prepectal carina is absent in the vast majority of these insects but a

few scarce tropical genera of each subfamily (except Khoikhoiinae) have the

character in plesiomorphic condition (Mason 1981).

13, Veins rl and r3 can be seen, as stubs in a few Cardiochiles (Mason

1981) species and a strong rl exists in Wesmaelella. The bend in IRs visible

in Cardiochiles and Microplitis tor example, occurs at the site of the

origin of the former vein rl.

19. The Y-shaped groove on metasomal tergum | (Fig. 20) is a charac-

teristic feature of Cardiochilinae, elsewhere in Ichneumonoidea characterizing

only the Braconinae in what appears to be a convergent development. The

Microgastrinae do not usually show such a structure but it is quite well de-

veloped in Miropotes, a very primitive microgastrine (Mason 1981], Fig. 24) and

traces can be seen in most Apantelini. The greatly narrowed tergite of

Miracinae obscures the Y-shaped groove but it can be traced in species with

larger tergites.

20-23. The structure of the metasomal tergum | in Cardiochilinae is

unique among Ichneumonoidea but has never been carefully described (Figs.

20, 21). The posterior half, more or less, is weakly sclerotized and undif-

ferentiated from side to side (except for the Y-shaped groove medially) and

has the spiracles situated laterally facing downward. The same relationship

of parts is found in many symphyta, e.g., Siricidae, Anaxyelidae, Orussidae,

58 Contrib. Amér.- Ent, Inst:, vol-20; 1983

Cephidae, Xyela and Megalodontes, but in no other Ichneumonoidea. In the

last group there is always (except in Cardiochilinae) a strongly sclerotized and

strongly differentiated median tergite and a pair of lightly sclerotized

laterotergites. :

The anterior half of the first metasomal tergum of Cardiochilinae shows

a normally differentiated and thickened tergite and thinner laterotergites,

these latter being continuous with the posterior parts of the tergum. The

sharp lateral ridges that delimit the tergite become lower and then merge,

toward the mid-length of the tergum, into the general surface. The tergal

development in Cardiochilinae seems to be intermediate between the undif-

ferentiated tergum of sawflies and the sharply defined tergite of typical

Ichneumonoidea. A very few highly specialized Cardiochilinae (Heteropteron,

Wesmaelella) have a completely defined mesal plate on tergum 1.

In Microgastrinae the lateral margins of the tergite | (Figs. 17, 18, 19)

are distinct from base to apex and the laterotergites exist as separate plates.

The unique feature here is that these laterotergites include the spiracles, a

condition found nowhere else in the Apocrita, although existing in

Xiphydriidae and Pamphiliidae among the Symphyta.

The lateral margins of tergite 1 are also complete in Miracinae and

Kheikhoiinae but the laterotergites have disappeared (Figs. 1, 9), leaving the

spiracles in membrane - again a unique condition for the Hymenoptera.

Nevertheless it is an open question whether the condition of the sides of

tergum | in these two subfamilies is a case of parallelism. I think that be-

cause the rest of the tergal structures are so very different in the two, they

have evolved in parallel.

The division of metasomal sternum | into an anterior, heavily sclero-

tized plate and a posterior, lightly sclerotized one is a basic feature of

Ichneumonoidea (Mason 1981). Nevertheless the "microgastri" do not clearly

show this division; the sclerotized portion of their first sternum being a

single rectangular plate that usually extends the full length of the segment

(Figs: 17-21); ,

If this plate be interpreted as an original condition, i.e, plesiomorphic,

then the rest of the Ichneumonoidea should be regarded, because of the

apmorphic divided first sternite, as a sister group to the microgastri, which

should have at least family rank. On the other hand, should the plate be

secondary, i.e., a product of the anterior plate alone, the posterior plates

having become desclerotized, or of the fused anterior and posterior plates,

then the structure will be apomorphic compared to its homologue in other

Ichneumonoidea. Some Microgastrinae have a harder looking median triangle

(apex caudad) that vaguely cuts off 2 softer looking posterolateral sclero-

tized areas (Fig. 17). This suggests that the large sternite may be compound.

For purposes of this study I have taken the view that the first sternite as

found in "microgastri" is a secondary fusion product. In either case, primary

or secondary, it is still a character of major importance.

REFERENCE

Mason, W.R.M. 1981. The polyphyletic nature of Apanteles Foerster

(Hymenoptera: Braconidae): a phylogeny and reclassification of Micro-

gastrinae. Mem. Ent. Soc. Can. 115, pp. 147.

BwWN

Mason: Khoikholinoo

oups

with apomorph.

—

TASLE, 1

Pe ga OR cares On sctllee bc aay ue Gnkenae ee Nene Ua en eres ne hg son era a ae oe?

Character ples. apo Notes : pany

. Flagellomeres-number variable constant

. Flagellomeres-number 16 12 a

. Antennal placodes irregular 2-ranked a

. Maxillary palpal

articles 6 1,2 fused r

. Maxillary palpal

articles : 5,6 fused a T

. Labial palpal 4 3 ar hor

articles 4 3 ant

. Clypeal margin convex straight to

concave

. Prepectal carina absent ie

. Transcutal groove absent hr

. Axillae horizontal vertical, hy

small

. Scutellar apical foveae absent present a

. Propodeum median absent present h

carina (areolate)

. Fore wing, rl, r3 absent hr

. Hind wing, rl absent ah ft

. Hind wing, r-m2 absent ho

. Hamuli, number 7-4 3 r

. Basitarsus 3, ventral present

ridge

. Tarsal claws, teeth absent hr

. Tergite M1, median present a oi

groove

. Tergite Ml, sides de- present a

limited basally only

. Tergite M1, sides present h

complete

. Tergite M1, latero- present a. A

tergite separated

. Tergite M1, latero- present absent ie

Notes: h - homeoplasy; a - autapomorph;

tergite

p - partial,

Total apomorphs

r — reductional apomorph;

meaning the apomorphic condition appears in some or most of

eae

M X

~ X

P Xx

Me xX

P P

Mee

P X

Mak

P X

P Xx

Meee

M X

P xX

omer

Me oe

_ Xx

Li oe

the more specialized species in the subfamily although the plesiomorphic

state seems to be the basic condition; C - Cardiochilinae;

K - Khoikhoiinae; M - Microgastrinae; X - Miracinae.

1983

Contrib. Amer. Ent. Inst., vol. 20,

"tkuuay “S$ ‘8

Stsueded “Ss ‘/ :auaweued pue SNYIBIP SPT[AS[OA ‘eTTe1IUSeH aTew ‘g‘/

"szoedse yesuop puke TeTOe} ‘pesy ‘osetuofuew “Ss ‘9‘G “uaWOopge ‘STsSueded

i ee

"§ ‘Z “Teue 1e] ‘eetuofuew eTUeS ‘|

*XxPU0YT JO WNSUOp ‘STSUeded “Ss ‘¢

é oy

Ook

SS os

"eHue1 Teseq ‘oaetuofuew

“Awoyzyeue etues “°g-| “SHTY

Mason: Khoikhoiinae

*uawopge ITSOUMOZ “y ‘G| °“S zOedSe [eSUuOp

pue yeToeR} “pesy “TSOUMOZ “yy ‘yl “E) “epPSnpetuios ~y ‘Z| <ezeTOS ~y

‘1, :uawopqe aTew Jo xode [eu.UaA ‘71 ‘1, °xXeuOoYy. JO WNSUOp ‘ezeTOS

"¥ “OL “Teva ze] “TseuMOZ “y “G6 “AwoyeUe BTOYYTOYY ~Gi-6 “SBT4

Contrib. Amer. Ent. Inst., vol. 20, 1983

Figs. 16-21. 16, wing of S. capensis. 1/7, Apanteles crassicornis

Prov., ventrolateral view of detached abdomen base to show apparent sub-

division of sternite 1. 18,19, Alphomelon sp., lateral and dorsolateral

views of base of abdomen. 20,21, Cardiochiles sp., base of abdomen: 20,

dorsal aspect, somewhat flattened, showing side plates; 21, dorsolateral

A REVIEW OF MYRMECOPHILIC-SYMPHILIC DIAPRIID WASPS IN THE

HOLARCTIC REALM, WITH DESCRIPTIONS OF NEW TAXA AND A

KEY TO GENERA (HYMENOPTERA: PROCTOTRUPOIDEA: DIAPRIIDAE)

Lars Huggert

Zoological Institute, Department of Systematics

S-223 62 Lund, Sweden

Lubomir Masner

Biosystematics Research Institute, Agriculture Canada

Ottawa, Canada KIA 0C6

Abstract

Myrmecophilic-symphilic members of twelve genera of the tribe Diapriini

(Diapriidae) in the Holarctic realm are compared and keyed. The following

new taxa are described: Labidopria noctivaga, n. sp. (Arizona), Plagiopria,

n. gen., with P. passerai, n. sp. (France, Italy), P. besucheti, n. sp.

(Turkey) and P. huberi, n. sp. (Iran), Townesella, n. gen., with

T. marjoriae, n. sp. (Arizona), and Trichopria myrmicae, n. sp. (New York).

The previously unknown male of Ecitovagus gibbus Masner is described. The

first known diapriid associates of ants of the genera Myrmica Latreille and

Plagiolepis Mayr are described; Plagiopria passerai is the first diaprid

positively reared from an ant (queen pupa). Available host records and

myrmecophilic habits of the wasps involved are discussed. A working hypoth-

esis to interpret the possible interrelationships between the ants and wasps

is presented.

INTRODUCTION

Initially, the aim of this paper was only to describe a few new taxa of

myrmecophilic Diapriini. However, as the work progressed it became obvious

that a critical review of all the genera involved and information accumulated

was desirable. As a result the myrmecophilic-symphylic members of twelve

genera (two of which are described here as new) known to occur in the

Holarctic realm were studied, compared and keyed. The present paper is

focused on highly integrated associates of ants, the symphiles as interpreted

by Masner (1977). Consequently, the non-symphilic associates such as the

synoekets or adventitious species of diapriid wasps are not considered. We

suspect that at least some of the frequent records of diapriine wasps in ant

nests (Kieffer 1916; Donisthorpe 1927) account for either the parasite's search

for dipterous hosts or for temporary shelter (e.g. hibernation).

Five symphilic genera are restricted to the Palearctic region, viz.,

Cordylocras Kozlov, Lepidopria Kieffer, Plagiopria, n. gen., Solenopsia

Wasmann, and Tetramopria Wasmann, and six symphilic genera are exclusively

Nearctic, viz., Auxopaedeutes Brues, Bruesopria Wing, Ecitovagus Masner,

Labidopria Wasmann, Myrmecopria Ashmead and Townesella, n. gen. All

these genera are either monotypic or contain only a few described species.

Only the large world-wide genus Trichopria Ashmead (s. 1.) with numerous

non-myrmecophilic species is represented in both the Palaearctic and Nearctic

regions by one species each positively associated with ants. Whereas the

64 Contrib. Amer. Ent. Inst., vol. 20, 1983

Palearctic genera seem to constitute a rather homogenous complex, the

Nearctic genera appear to be sharply divided into two groups. Auxopaedeutes

and Bruesopria are closely related to the Palearctic genera Lepidopria

and Solenopsia because of non-compressed tarsi and an abrupt antennal

clava in the female. Ecitovagus, Labidopria, Myrmecopria, and

Townesella on the other hand, show distinct ties to several Neotropical

ecitophilic genera (Masner 1977) because of compressed tarsi and a non-

abrupt multisegmented antennal clava in the female. We believe that all

symphilic members of the above genera can be derived from different

species-groups of the polytypic genus Trichopria s. 1. In our opinion

social integration is the driving force in this evolution.

Three groups of ants are involved as the main hosts of myrmecophilic

diapriines, viz. the Myrmicinae, Formicinae and Dorylinae. Among the

Myrmicinae the genera Solenopsis Westwood, Tetramorium Mayr, and possibly

also Myrmica Latreille have integrated diapriines. Formicine hosts include the

genera Formica Linné, Camponotus Mayr, Lasius Fabricius and Plagiolepis

Mayr. However, only Plagiolepis is proven here to be a real host while the

other genera may be just circumstantial records (e.g. numerous species of

Trichopria and Basalys (=Loxotropa acut.)( Kieffer 1916). The doryline ants

of the tribe Ecitonini in the New World such as the genera Labidus Jurine,

Neivamyrmex Borgmeier, Nomamyrmex Borgmeier and Eciton Latreille host a

large number of highly specialized diapriine symphiles (Borgmeier 1939;

Ferriere 1929; Loiacono 1981). This specialization is highlighted by phenomena

such as mimicry, alectomy, dual phase of adult life, complete social integra-

tion into ant communities, etc.

We presume that the Trichopria-like ancestors of extant ecitophilic

symphiles gradually shifted from parasitizing dipterous hosts in general to

parasitizing ecitonine ants. The intermediate step in this presumed sequence

of hosts seems to be the numerous synoeketic Diptera living in the refuse

depots and bivouacs of various army ants (Wilson 1972). The presumed life

cycle of a diapriine symphile consists of three phases, viz. postembryonic,

dispersal and social (Fig. 1). Following completion of postembryonic develop-

ment in the ant brood, the fully winged wasps (F,M) emerge from the ant

nest and are subsequently engaged in nuptial and dispersal flights. These

flights are largely nocturnal as documented by frequent interceptions of wasps

in UV light traps. During the peak of this slight alectomy evidently begins,

as individuals joining the ants are already dealated and the individuals at-

tracted to light traps show various degrees of dealation. In the third (social)

phase the wasps assume new functions such as taking part in raids and other

activities as members of the ant society. Sometimes during this phase the

wasps parasitize ant brood. Most species seem to have males only faculta-

tively or not at all. To date only Ecitovagus gibbus Masner is shown here

to have dealated males in the third phase of the life cycle.

The known bionomics of the non-ecitophilic genera of myrmecophilic

Diapriinae present a very interesting enigma. Based on the classical works

on myrmecophilism by Erich Wasmann at the turn of this century, members

of these genera are generally believed to be true myrmecophiles (symphiles),

possibly parasites of ants. This belief was based primarily on a high degree

of integration of these wasps with the host ants, the adaptations being both

Huggert & Masner: Diapriid wasps 65

morphological and behavioural. However, until now there has been no posi-

tive rearing record of diapriines parasitizing any stages of host ants (Wing

1951). Plagiopria passerai, n. gen. et n. sp. is the first diapriine reared

from a host ant. Luc Passera (Toulouse in lit.) reports the above wasp

as reared from the ant Plagiolepis pygmaea (Latreille): "Ils (5 F) sont

tous sortis de coccons de reine de cette fourmi au laboratoire". On the con-

trary, there are several well documented host records of members of genera

having myrmecophilic species reared from puparia of various Tachinidae (Dip-

tera). Both sexes of Tetramopria aurocincta Wasmann were reared from a

puparium of Compsilura concinnata Meigen, a primary parasite of the fall

webworm, Hyphantria cunea (Drury), in Hungary (Szelényi 1957). Later, the

same species of wasp was reared again under similar circumstances in

Czechoslovakia (Masner in Jasic 1964). The senior author examined a series

of Tetramopria aurocincta labelled "Obtenu de graines Phlox nains" in

[MNH], Geneve. An interesting case of simultaneous parasitism by

Lepidopria pedestris Kieffer and Trichopriainquilina Kieffer on a tach-

inid puparium in an adult may beetle, Melolonthamelolontha Linne is known

to us from Yugoslavia (specimens in Nat. Museum, Praha). Bures (1916) des-

cribed Lepidopria aberrans from a puparium of the tachinid Cryptomeigenia

theutis (Walker) found in a dead adult june beetle, Phyllophaga inversa

(Horn), in the USA; examination of the unique type (USNM, on slide) of

Brues' species indicates that it probably belongs to Bruesopria Wing. Sever-

al specimens of Auxopaedeutes sp. reared from cocoons of a diprionid sawfly

(? parasitized by a tachinid) were studied by the junior author (USNM,

Washington).

The above conflicting evidence poses an interesting biological problem.

What is the role of these highly integrated wasps in ant societies and what

are their true hosts? Diapriines are, except for few problematic records,

primary parasites of various cyclorrhaphous Diptera. Therefore, we are in-

clined to believe that the female wasps, while pursuing the potential dipterous

hosts in the soil, may enter ant nests. Gradually, during this process, the

wasps have become increasingly associated and integrated with the ants.

Initially, this gave a wasp a choice of parasitizing either dipterous puparia

or the similarly shaped ant brood. The shift from dipterous to formicid hosts

also has determined the degree of integration which in turn controls the

mechanism of morphological and behavioural specialization. The twelve

myrmecophilic-symphilic genera discussed in this paper display various degrees

of social integration and specialization.

This paper deals primarily with new taxa, however, for data concerning

type-species, synonymies, etc. of the genera dealt with in the following key,

see Muesebeck and Walkley (1956).

ABBREVIATIONS OF MUSEUMS

CNC - Canadian National Collection of Insects, Ottawa, Canada.

FMCH - Field Museum, Chicago, USA.

HUGG - Collection Lars Huggert, Lund, Sweden.

LPT - Collection Luc Passera, Toulouse, France.

MCV - Museo Civico di Storia Naturale, Verona, Italy.

66 Contribs Amer. Ent. inst., vol. 20, 1983

MNH - Musee d'Histoire Naturelle, Geneve, Switzerland.

UAT - University of Arizona, Tucson, USA.

USNM - United States National Museum of Natural History, Washington, USA.

KEY TO GENERA OF MYRMECOPHILIC-SYMPHILIC

DIAPRIINAE IN HOLARCTIC REALM

1. Anterior margin of T2 (between corners) at most as wide as petiole

(Figs. 2, 3, 17, 18); tarsi strongly compressed laterally, with tarsomeres

much higher than wide; female antenna with undifferentiated 7-8 seg-

mented clava (Figs. 20, 21); anterior margin of S2 apparently without

longitudinal groove-like invaginations; rufous or xanthic forms associated

With ecltenine,aots in. Now Would. 8 ee ces ee ee ee ee eee a

Anterior margin of T2 (between corners) moderately to distinctly wider

than petiole (Figs. 4-7, 30); tarsi cylindrical, with tarsomeres about as

high as wide (except in Plagiopria huberi); female antenna usually

with abrupt 2-4 segmented clava (Figs. 23, 24, 39); anterior margin of

S2 usually with two more or less distinct groove-like invaginations;

associated with myrmicine and formicine ants in both Old and New

RS a a eg 5

2. Head and mesosoma including tegulae mat, granulose-rugulose; meso-

scutum with high hump anteromedially; propodeum much protruded

posteromedially (Fig. 3); A3 in female 3x longer than A2.

Only one species, E. gibbus Masner associated with Neivamyrmex

nigrescens (Cresson).in Arizona (M, FF)... . . 0... Ecitovagus Masner

Head and mesosoma at least partly smooth and shining, with tegulae

smooth; mesoscutum not humped anteromedially; propodeum not pro-

truded bosteramedially (Figs. 2, 17, 18); A3 in female as long as or

only slightly longer than A2 (Figs. a oS ee 3

3. Head globose, slightly wider than long, with all ocelli inside ocular zone

(Fig. 15); petiole with distinct tooth anteromedially (Fig. 13); female

antenna 12-segmented, with apical segment very small (Fig. 21); mid

coxae with dense tuft of hairs interoapically (only in L. noctivaga,

n. sp.). Several undescribed Neotropical species; L. longicornis

Wasmann (Brazil) and L. costaricana Borgmeier (Costa Rica) both

with Eciton praedator F. Smith. Only L. noctivaga, n. sp. in

Avizona (Fy oo a ee ees Labidopria Wasmann

Head slightly to distinctly elongate, with posterior ocelli clearly outside

ocular zone (Fig. 16); petiole without tooth anteromedially; female

antenna ll-segmented, with apical segment enlarged (Fig. 20); mid

coxae without dense tuft of hairs interoapically ............- 4

4. Head in dorsal view with temples subparallel, broadest at occiput; head

with distinct irregular sculpture and with transverse sinuate ridge on

frons; pronotal shoulders rounded; dorsal apices of tibiae strongly pro-

jecting, tooth-like. Only M. mellea (Ashmead) associated with

Neivamyrmex opacithorax (Emery) and N. carolinense (Emery) in

Southeastern USA. (My. FURL Ser has Myrmecopria Ashmead

Head in dorsal view with temples converging towards occiput, broadest

across eyes (Fig. 16); head smooth and shining, without transverse

M = Male; F = Female.

Huggert & Masner: Diapriid wasps 67

ridge on frons (Fig. 14); pronotal shoulders angularly protruding (Figs.

16, 19); tibial apices not projecting. Only T. marjoriae,n. sp. in

AupOna UP bas 2. cases BEE Clas Townesella, n. gen.

5. Head in dorsal view posteromedially strongly tooth-like projecting, be-

tween this projection and bluntly backwards projecting temples occiput

deeply excavate (Figs. 25, 26); pronotum with horn-like projecting

shoulders (Fig. 27). Only c. mirabilis Kozlov in Azerbaijan and

Ciineas A) a ees uns ce a Ba ae ee ee Cordylocras Kozlov

Head in dorsal view more or less globose, without projections or depres-

sions; , prenotal.shoulders not projecting... < . 408 ci4eee bas eee Gus 6

6. Petiole in lateral view humped dorsally, distinctly elevated above level of

anterior mare tor 12 (Ming (921) a ak ee 7

Petiole in lateral veiw not humped, cylindrical, not elevated above level of

anterior margin of T2 (Figs. 8, 12)

7. Petiole in dorsal view distinctly broadened posteriorly (Fig. 4); propodeum

anteromedially with sharp triangular keel (Fig. 9); female antenna lII-

segmented, with 2-segmented clava; genae ventrally with thick brush

of short setae. Only S. imitatrix Wasmann, associated with

Solenopsis fugax Latreille in Europe. (M, F) . Solenopsia Wasmann

Petiole in dorsal view not broadened posteriorly (Figs. 5, 6); propodeum

anteromedially without elevated keel (Figs. 10, 11); female antenna ll-

or 12-segmented, with 3- or 4-segmented clave; genae ventrally at most

Wil) Spatse silvery Balte 4 i sighed des ee Sciuele Gee acer 8

8. Propodeum deeply excavated posteriorly with flanges elevated, propodeum

very short medially, with no median keel (Fig. 5); petiole with hump

directed backwards concealing anterior margin of T2 at meson (Figs.

5, 11); female antenna ll-segmented, with abrupt 3-segmented clava.

Only B. americana (Brues) and B. seeversi Wing, associated with

Solenopsis molesta (Say) in North America. (F) . . Bruesopria Wing

Propodeum not excavate posteriorly, without flanges, longer medially, with

distinct median keel (Fig. 6); petiole with hump directed upwards not

concealing anterior margin of T2 at meson (Figs. 6, 10); female an-

tenna |2-segmented, with distinct 4-segmented clava. Only

L. pedestris Kieffer, associated with Solenopsis fugax Latreille in

Puree. (RU i ee Lepidopria Kieffer

9. Propodeum deeply excavated dorsomedially, with prominent flanges at

sides (ieee Be 10

Propodeum at most slightly concave posteromedially, with no specialized

flanges at sides (hie 2 45) he ew pe ee a a

10. Hind tibia strongly compressed laterally, with sharp blade-like edges in

proximal ventral halves (Figs. 32, 36); petiole more or less glabrous

and smooth dorsally, with short to long hairs posterolaterally (Figs. 8,

30, 35, 37); scape reticulate; male A5-AI4 distinctly elongate (Fig.

33). P.passerai, n. sp., associated with Plagiolepis pygmaea

(Latreille) in S. Europe, P. besucheti,n. sp. (Turkey) and P. huberi,

As GO d emai Wed Rhye digs (ack ehivene ae ee a Plagiopria, n. gen.

68 Contrib. Amer. Ent. Inst., vol. 20, 1983

Hind tibia not compressed laterally, with no sharp edges ventrally;

petiole dorsally with minute scales and dense long hairs (Fig. 7);

scape smooth; male A5-Al4 penicillate. Six species in N. America,

A. sodalis Brues and B. lyriformis Brues associated with Solenopsis

mOolee on Aaa) le os. oak es eae ee Auxopaedeutes Brues

11. Head in dorsal view sub-hexagonal, with sides almost straight; tufts on

pronotum and temples golden yellow; eyes remarkably small. Two

species, T. aurocincta Kieffer and T. cincticollis Wasmann asso-

ciated with Tetramorium caespitum (Linné) in Parope tM, FF) ero

pete rah ue Te a 4 ee a: Tetramopria Wasmann

Head in dorsal view usually globose, with sides rounded; tufts on pronotum

and temples whitish; eyes usually larger. Several species reported as

associated with ants in Europe (Kieffer 1916; Pschorn-Walcher 1957,

Dessart 1975), T. inquilina (Kieffer) with Solenopsis fugax

Latreille in Europe and Tf. myrmicae,n. sp. with Myrmica americana

Weber in iN: Ameritg. (MR) ecg yh ea, Trichopria Ashmead

1. Labidopria noctivaga, n. sp. (Figs. 13, 15, 17, 21)

Female: Length about 1.9 mm. Honey colored, with metasoma slightly

darker, eyes and ocelli blackish and all sutures dark brown; wings hyaline;

body predominantly smooth and shining, with scattered long semidecumbent

setae.

Head (36:42) smooth and shining; in dorsal view eyes distinctly longer than

strongly receding and rounded temples (Fig. 15); temples without pilosity; ocelli

inside ocular zone; POL:LOL:00L = 6:5:10; occipital collar unusually narrow; eyes

in lateral view higher than long (23:18), with 12 and 8 ommatidia respectively;

rather raspberry-like; malar space shorter than length of eyes (13:18) (Fig.

13); mouth small, mandibles bidentate, with lower tooth sharp, considerably

longer than upper tooth; palpi short, formula 3-2; hypostomal carina fine, not

particularly projecting; antenna without differentiated clava (Fig. 21), A4-Al2

subquadratic, closely approximated; Al2 unusually small, shorter than preced-

ing segment; apical rims of A4-A11 with irregular sprockets; flagellomeres

with delicate granular microsculpture except for smooth spot on Al apico-

ventrally (between flanges); scape reticulate, seen dorsally distinctly com-

pressed basally, gradually incrassate towards apex, with rather large apical

flange housing pedicel in flexed position; antennomeres in relative proportions

SO 1Os Os PerdOG; To GUS Te Paes PAO See 7 Or 7 SOs OE 6.527.

Mesosoma as high as wide; mesoscutum wider than long (19:12); pronotal

shoulders smooth and glabrous, slightly angularly projecting; small tuft of

yellow hairs below pronotal shoulders; scutellar pit shallow, inconspicuous;

axillae and scutellar shield subcircular, lateral margins of shield delicate;

propodeum short, shorter than scutellar shield (13:17), with no central keel

or plicae and with posterolateral corners not projecting, with minute semi-

transparent flanges (Fig. 17); propleura with sparse whitish pilosity; anterior

margin of pronotum with no tuft; metapleura and sides of propodeum with

only few scattered long setae; metapleura smooth, with deep depression

anterodorsally connecting to depression between metanotum and propodeum;

metapieural carina not clearly defined; mid coxae apically with dense tuft of

yellowish hairs interoventrally; hind coxae unusually long (29:11),

Huggert & Masner: Diapriid wasps 69

subcylindrical; marginal cilia of fore wing 1/5 of wing width; fore wing sur-

passing metasoma by half of its length.

Petiole (25:13) in dorsal view subcylindrical, distinctly constricted at apex

widest in posterior third, with fine granular sculpture dorsally and rougher

sculpture laterally; in lateral view petiole with prominent tooth antero-

ventrally (Fig. 13); rest of metasoma short, only slightly longer than wide

(67:45).

Male: Unknown.

Biology: Unknown; caught in UV light trap in semidesert habitat; pre-

sumably associated with army ants.

Variability: The specimens are very uniform in all respects and no sub-

Stantial variation was encountered.

Distribution: Arizona.

Type material: Holotype: Female, USA, Arizona, Pima Co., 8 mi. N.

Vail, Aug. 30, 1962 (F. Werner & W. L. Nutting) [USNM]. Paratypes :

4 F, USA, Arizona, Brown Co., Baboquivari Mts., Aug. 4, 1962 (F. Werner &

P. Johnson); USA, Arizona, Pena Blanca, Sta. Cruz Co., July 10, 1961 (F. Werner

& W. L. Nutting); USA, Arizona, Tumacacori, Santa Cruz Co. April 10, 1975

(G. Ginelly) [CNC No. 17551, UAT, HUGG].

Etymology: The name of this new species refers to its presumed nocturnal

dispersal flight preceeding the wing autotomy and presumed association with

some species of army ants.

Remarks: This new species is placed in Labidopria with some reservation.

It differs from all described as well as several undescribed species of the

genus in having a strong tooth situated anteroventrally on the petiole and

tufts of hairs on the mid coxae. Furthermore, the new species differs from

L. longicornis Wasmann (type-species of Labidopria) and L. costaricana

Borgmeier by having a minute propodeal flange, narrow occipital collar, as

well as by having a different shape of petiole. L. noctivaga is the northern-

most species of Labidopria, and at the same time the first record of the

genus from America north of Mexico.

TOWNESELLA, new genus

(Figs. 14, 16, 18, 19, 20)

Female: Body honey yellow, predominantly smooth and shining, with

scattered long semidecumbent setae; head in dorsal view slightly longer than

wide, subhexagonal, with temples straight but somewhat convergent and with

antennal shelf small; at least posterior ocelli outside ocular zone (Fig. 16);

eyes large, with raspberry ommatidia; mouth small, mandibles bidentate, with

lower tooth sharp and much longer than upper tooth; palpi short, with formula

3-2; hypostomal carina fine, not particularly projecting; clypeus not delimited

above; temples behind without pilosity but with small area of granular sculp-

ture appearing as minute sharp points (Fig. 19); antenna (Fig. 20) ll-segmented,

without differentiated clava, with antennomeres A4-Al0 subquadratic, closely

approximated; all considerably longer than preceeding segments; apical rims

70 Contrib. Amer. Ent. Inst.,-voly 20, 1983

of A4-Al0 with irregular sprockets; scape reticulate, somewhat compressed

basally (dorsal view), with minute apical flange.

Mesosoma slightly flattened dorsally and in dorsal view rather narrow

(Figs. 14, 16); pronotal shoulders distinctly prominent, rounded, with delicate

granular microsculpture similar to that on temples; notauli absent, mesoscutum

with setae in two submedian rows; scutellum with large subrectangular, mod-

erately deep pit, with scutellar shield subcircular with setae in two submedian

rows; scutellum with large subrectangular, moderately deep pit, scutellar

shield subcircular with delicate lateral margins; metanotum unarmed medially;

propodeum short, with sharp prominent median keel and with prominent blade-

like flanges posterolaterally (Fig. 18); sides of propodeum and metapleura

glabrous, with only few scattered long setae; metapleural carina distinctly de-

fined, metapleuron with vertical groove anteroventrally; mesopleuron with dis~

tinct oblique line; fore wing long, with rather long marginal cilia; submarginal

vein short, slightly exceeding basal 1/4 and closely paralleling front margin

of wing; basal vein not indicated; hind wing narrow, with submarginal vein

reduced to rudiment at base; legs long and strong, with tarsi distinctly com-

pressed, much higher than wide; hind coxa rather large, broadest in basal

third, gradually tapering to apex.

Petiole (Fig. 18) in dorsal view subcampanulate, only slightly longer than

wide, distinctly constricted anteriorly and with posterior margin sharp, slight-

ly concave; rest of metasoma short, only slightly longer than wide; anterior

margin of T2 straight.

Male: Unknown.

Type-species: Townesella marjoriae, n. sp. (described below).

Etymology: This new genus is named in honor of Henry K. Townes at the

occasion of his life jubilee. The suffix -ella is a diminutive in reference

to the small size of this wasp. The gender is feminine.

Remarks: This new genus appears to be related to other predominantly

smooth-bodied members of known genera of ecitophilic Diapriini (e.g.

Asolenopsia Kieffer, EFuplacopria Ferriere, Labidopria Wasmann and

Neivapria Borgmeier) by the following peculiar characters: Female antennae

with secondarily nondifferentiated clava, tarsi strongly compressed laterally,

body uniformly honey-colored, and petiole more or less specialized.

Townesella is closely related to Asolenopsia because of having an ll-segmented

antenna with the apical segment clearly longer than the preceeding ones.

However, this new genus differs markedly in the development of body pilosity

in that Asolenopsia has the temples and the anterior margin of the pronotum

with dense tufts, and the propleura, metapleura as well as the dorsum of the

petiole hairy. Townesella, on the other hand, has no specialized pilosity in

these areas, except for the usual scattered semidecumbent setae over all the

body and a dense brush of hairs ventrally on the petiole. The closest relative

of Townesella is probably Myrmecopria Ashmead because of the similar shape

of the petiole, the elongate head, as well as the general structure of the

propodeum. For differences between these two genera, see the key.

Huggert & Masner: Diapriid wasps cs

2. Townesella marjoriae, n. sp. (Figs. 14, 16, 18, 19, 20)

Female: Length 1.7 mm. Honey colored, dorsal part of head and body

of metasoma darker and petiole, mesosoma, legs and antennae lighter; eyes

and ocelli blackish and all sutures dark brown; wings hyaline.

Head (38:37) smooth and shiny, in dorsal view with eyes subequal to tem-

ples; POL:LOL:00L = 4:5:12.5 (Fig. 16); eyes in lateral view higher than long

(19:14), with about 12 by 8 ommatidia respectively; malar space shorter than

length of eye (11:14) (Fig. 14); flagellomeres (Fig. 20) with delicate granular

microsculpture except for smooth spot on Al apicoventrally between flanges;

Al reticulate; antennomeres in relative proportions 28:9.5; 9:6; 9:6; 6:7;

6:25; O:Bs- 6 B29 Tie cIOe, 7293 -16:9:

Mesosoma somewhat higher than wide (42:38); mesoscutum as long as

wide; pronotal shoulders without tuft of hairs but with mat granular micro-

sculpture; suture between axillae and scutellar shield thickened, finely crenu-

late; axillae and scutellar shield partially with delicate longitudinal micro-

sculpture; propodeum short, shorter than scutellar shield (19:13), steeply slop-

ing, with sharp median keel anteriorly, deeply excavate between long flanges,

with granular sculpture; flanges of propodeum semitransparent, finely longi-

tudinally sculptured, in dorsal view divering apically (Figs. 14, 18); metapleuron

evenly and coarsely reticulate; marginal cilia of fore wing 1/7 of wing width,

surpassing metasoma by half its length; hind coxa (20:14) basally with small

patch of coarse sculpturing.

Petiole (16.5:15) entirely roughly reticulate, with sharp posterior margin

and with brush of short dense setae ventrally; T2 large, occupying almost

entire body of metasoma.

Male: Unknown.

Biology: Unknown; caught in UV light trap in semidesert habitat; pre-

sumably associated with army ants.

Variability: The specimens are very uniform in all respects and no

substantial variation was encountered.

Distribution: Arizona.

_ Type material: Hologype : Female, USA, Arizona, Molino Basin, Sta.

Catalina Mts., August 2, 1978 (C. A. Olson & M. W. Hetz) [USNM];

Paratype : 10 F, with same data as in holotype but caught on July 7, 1962

(F. Werner); USA, Arizona, Pima Co., August 30, 1962 (F. Werner & W. L.

Nutting); USA, Arizona, Brown Co. Baboquivari Mts., August 4, 1962

(F. Werner & P. Johnson); USA, Arizona, Santa Cruz Co. 10 mi NW Nogales,

September 4, 1968 (T. R. Haig) [UAT, USNM, CNC no. 17552, HUGG].

Etymology: This new species is named in honor of Dr. Marjorie Townes,

a life time companion of Henry Townes.

3. Ecitovagus gibbus Masner 1977 (Figs. 28, 29)

The male of this species was not previously Known. It differs from the

female in some minor characters.

72 Contrib. Amer. Ent. Inetepwol: 20: 1983

Male: Length about 2.8 mm. Color and general shape of body as in fe-

male; sculpture of mesosoma and particularly of head slightly finer than in

femaie; antennal segments (Fig. 28) in proportions 59:13; 12:9; 22:12; 19:15;

Heh isisy 103s 107,5* t0:12.97 1012.5: 95:12: OZ: 9:12: 9:93, AS and. A4 much

modified (Fig. 29); A3 strongly horizontally dilated apically with anteroventral

surface flattened; A4 also strongly dilated but vertically, triangular, with ex-

terior surface concave; A5-Al3 cup-shaped, proximal ones distinctly asym-

metrical exteriorly; Al4 much smaller than Al3; Al sculptured as in female;

antenna bristly; setae on body, especially on metasoma, distinctly longer,

about twice as long as in female; female petiole only with some scattered

setae except for anteromedial triangular tuft of adpressed golden hairs; male

petiole covered all over with dense yellowish setae, except for sparsely setose

anteromedial slightly depressed area; metasoma shorter than in female with

more obtuse apex.

Biology: The individuals from Cochise Stronghold are the first record of

E. gibbus caught in light traps. The alate individuals disperse probably dur-

ing both day and night, the dealate individuals are probably strictly nocturnal

in habits.

Additional material: Female, U.S.A., Arizona, Cochise Stronghold, Dragon

Mts., August 16, 1962, Light trap (F. Werner) [Hugg]; 2 M, as above but

caught August 21-24, 1969 and August 22-27, 1970, light trap (R. J. Shaw)

[CNC and UAT]; dealated F, Arizona, Cochise Co., Chiricahua Mts., South-

western Research Station nr. Portal, 5400 ft., August 2, 1977, taken from raid

column of Neivamyrmex nigrescens in night (L. Masner) [CNC]; alate F as

above but caught by sweeping plants in daytime on August 3 (L. Masner)

[CNC]; dealated F M, Arizona, Cochise Co., Chiricahua Mts., Cave Creek

Canyon, SW Research Station nr. Portal, 5500 ft., August 13, 1965, taken from

night raid column of Neivamyrymex sp. CH. §. Dybas) [M, CNC and F, FMCH ].

PLAGIOPRIA, new genus

(Pigs, 8, 223: 24,30; 338)

General habitus of Ashmeadopria Kieffer (sensu Sundholm 1960), however

differing from the latter in the following characters.

Female: Head more or less globose with antennal shelf prominent; ocelli

small, OOL at least 3 x as wide as POL; head clothed with minute hairs but

no long setae; head smooth and shiny with some fine longitudinal rugulosity

laterally and anteriorly on antennal shelf (Fig. 31); in P. huberi this sculp-

ture is spread over most of head as peculiar, very open, finely impressed

foveolate reticulation; mouth very narrow; mandibles small, bidentate; palpi

short, formula apparently 5-2 (at least in M of P. passerai (Fig. 34); an-

tenna 12-~segmented, with moderate to indistinct 3-segmented clava (Figs.

22-24); A3 remarkably elongate (in two species), constricted basally, almost

as long as A4-A6 combined, but in P. huberi only slightly longer than A4;

A4-A9 globuliform, constricted basally; scape entirely with fine reticulation,

which is pronounced in P. huberi and here also confined to pedicel; flagel-

lum clothed with rather long bushy, dense pilosity.

Huggert & Masner: Diapriid wasps 73

Prothorax strongly developed at shoulders, almost rectangular anteriorly

but in P. huberi of more usual Trichopria-shape; cervical part with massive

collar of very dense short pilosity (only as two spots in P. huberi); meso-

scutum flat, almost triangular, glabrous but in P. huberi of a usual

Trichopria-shape and whole mesosoma with finely regulose sculpture; median

part of mesoscutum slightly elevated with shallow median depression; pro-

podeum (Figs. 8, 30) deeply excavate dorsomedially, glabrous in this area;

plicae elevated, blade-like, considerably bent inwards and projecting posterior-

ly into semi-transparent flanges; median keel reduced to a point (in P. huberi

virtually nonexistent) situated in extreme anterior part of propodeum; hairs

on sides of propodeum minute and sparse, not obscuring propodeal spiracles;

fore wing with only sparse hairs on basal third, costal cell very narrow; hind

wing with incomplete submarginal vein; legs peculiar (Figs. 32, 36), femora

gradually incrassate, not abruptly neck-like constricted basally (lateral view),

but strongly compressed laterally in basal third with sharp blade-like edges

in basal half or more; row of bristles on hind tibiae pronounced (except in

P. huberi); tarsi more or less compressed. |

Petiole smooth and shiny dorsally, not humped, more or less constricted

posteriorly (not obvious in P. huberi); anterior margin of T2 entire, only

slightly wider than petiole; T2 and following tergites with dense adpressed

to scattered longer hairs; S2 (ventral view) with two longitudinal deep and

broad carinated furrows in basal half.

Male: Similar to female but antenna 14-segmented, flagellar segments

each with whorl of errect bristles and A4 not modified (Fig. 33).

Type-species: Plagiopria passerai, n. sp. (described below).

Etymology: The prefix Plagio- (meaning to steal) is derived from the

name of the host ant, Plagiolepis pygmaea ; the suffix -pria meaning the

wasp. The gender is feminine.

Remarks: Plagiopria is a highly specialized member of the Trichopria-

complex in the tribe Diapriini. From Trichopria s. 1. it is distinguished

by the following peculiar character states: the sculpture of Al, the pilosity

of flagellum and the shape of A4-A9; the structure of propodeum and petiole;

the shape of legs, hind tibia in particular; the deep grooves on S2.

Plagiopria passerai is the first known diapriine associate of the ant genus

Plagiolepis Mayr (Formicidae).

There are three species of Plagiopria known to us, of which P.

passerai and P. besucheti seem to form one group and P. huberi another

group (see the key). P. huberi is certainly the most aberrant species, how-

ever, creation of an independent genus for it at this moment does not seem

justified because there may be more as yet unknown species in

considering the extensive distribution of the ants of the tribe Plagiolepini,

mainly in the Old World tropics (Brown 1973, personal communication).

KEY TO THE KNOWN SPECIES OF PLAGIOPRIA

1. Head and mesosoma predominantly sculptured; Al and A2 with pronounced

reticulation (Fig. 22); A3 only 1.7 x as long as wide; flagellum broadest

74 Contrib. Amer. Ent. Inst., vol. 20, 1983

proximally, clava inconspicuous and all antennal segments irregular in

outline; prothoracic shoulders not conspicuously angular; propodeum

without median carina; hind tibiae much narrowed in basal half, com-

pressed (Fig. 36); metasoma with scattered short hairs (Fig. 37);

yellowish brown (ran)e(F) 2 2. « ucec.. 6. Plagidpria huberi, nie sp.

Head (except for antennal shelf) and mesosoma smooth; only Al more or

less reticulate (Figs. 23, 24); A3 at least 2.5 x as long as wide;

flagellum broadest anteriorly, clava rather distinct and antennal seg-

ments smooth in outline; prothoracic shoulders angular; propodeum with

hump-like median carina (Figs. 8, 30); hind tibiae wide, subparallel

with short neck in basal sixth (Fig. 32) metasoma with hairs as in

Pigg 220. 37; Chestnt Drown 2 a eli eee os tee Be ee 2

2. T2 with dense short pilosity, laterally to lateroanteriorly directed apically

(Fig. 30); petiole with long hairs laterally; scape with fine but distinct

reticulation (Fig. 23); exterior margin of mid tibiae slightly sinuate;

pronotum in dorsal view with several hairs on each side close to scutal

suture (France, Italy). (M, F) ..... 4.Plagiopria passerai, n. sp.

Metasoma with only scattered long setae (Fig. 35); petiole with short

hairs laterally; scape delicately longitudinally reticulate (Fig. 24); ex-

terior margin of mid tibiae more sinuate; pronotum in dorsal view with

only two hairs on each side at scutal suture (Turkey). (F) .......

De ee ee ee 3.Plagiopria besucheti, n. sp.

4. Plagiopria passerai, n. sp. (Figs. 8, 23, 30-34, 38)

Female: Length about 2.0 mm. Color chestnut brown with sutures and

edges darker, as head and metasoma; wings slightly yellowish.

Head in dorsal view globose (Fig. 30) (34:35) with rather conspicuous

ridge between antennal sockets; head with rather dense minute hairs and

pilosity on temples rather sparse; eyes small, shorter than temples, glabrous;

ocelli close together, POL:LOL:00L = 4:4:15; head in lateral view (Fig. 31)

slightly longer than high (38:33), space between front orbit and apex of an-

tennal shelf with fine longitudinal rugulosity; eyes subcircular, subequal to

malar space; antenna (Fig. 23) with segments in proportions 35:7; 6:6; 13.5;

9:0; 9:0;° 5:6; °9:6.5; 5:6.5: 5.5:7; 9:8; 10:8:5; 1077; AlZ conspicuously narrower

than All and reticulation on scape pronounced; palpi very small, maxillary

palpus with very short apical segment.

Mesoscutum distinctly wider than long (32:45), with setae placed as in

Fig. 30; scutellar pit shallow, transverse, scutellar shield almost parallel-sided

with median carina hardly indicated and lateral margins fine; propodeum

rather long and narrow with some longitudinal wrinkles on projecting semi-

hyaline flanges (Figs. 8, 30); fore wing (Fig. 38) sparsely hairy on disc in

proximal third, with about 12 short bristles on submarginalis + marginalis;

marginal cilia slightly more than 1/8 of wing width; hind wing with cilia about

half as long as wing width; tarsi short, slightly compressed; femora basally

at constricted part with short, narrow, sharp edges, especially on ventral side

(Fig. 32); exterior margin of mid tibia slightly sinuate; hind tibiae wide, sub-

parallel with short neck in basal sixth and sharp interior edge medially.

Petiole (18:12) slightly excavate anterodorsally, with tufts of long hairs

laterally; body of metasoma about 1.7 x as long as wide; T2 (32:23) covered

Huggert & Masner: Diapriid wasps 15

by dense minute hairs, in apical fifth directed laterally to anterolaterally

(Fig. 30); last tergite triangular, pointed.

Male: Length about 2.0 mm. Color as in female but head and metasoma

slightly darker than in female.

Head as in female but antennal shelf smooth and less pronounced; eyes

somewhat larger; antenna (Fig. 33) with segments in proportions 33:6; 6:5;

16:45 14255: 18:55 92:36 12261225 12:5; 12:5; 12253 1225; 11:5; 164s AS not. modi-

fied; antenna slightly longer than body; flagellar segments with whorl of

erect bristles, which are clearly shorter than length of segments.

Mesoscutum as in female but laterally bent propodeal flanges slightly

wider apart; wings as in female but legs more slender with less pronounced

sharp edges and appearing less compressed.

Petiole and metasoma subequal to that of female but pilosity of T2 sparser

and two-directional pilosity apically on T2 less expressed.

Biology: The specimens from Montricoux (France) were reared from

pupae of queens of Plagiolepis pygmaea (Latr.) (Formicidae) (L. Passera,

in dies. |

Variability: In the material studied no variation of any importance

could be encountered.

Distribution: S and E France, northern half of Italy.

Type material: Holotype: Female, France (T. & G.) Montricoux,Sep~

tember ll, 1973 (L. Passera) [CNC No. 15596]. Paratypes: 5 F, 2M. 1F

with same data as holotype except the date, September 2 [CNC]; 2 F with

same data as first paratype [Hugg and LPT]; 1 F, France, Hte. Savoie, Grande

Saleve, April 26, 'Hymenoptere chez fourmis" [MHN]; 1 F, Italy, Veneto,

Monte Berici, February 20, 1972 (R. Pace) [MCV]; 1 M, Italy, Umbria, Monte

Peglia, Marsciano, August 9, 1979 [Hugg]; | M, Puglia, Monte Gargano, Foresta

Umbra, August 8, 1979 (both L. Huggert) [Hugg].

Etymology: The new species is named in honor of Dr. Luc Passera (Lab-

oratoire d'entomologie, Universite Paul Sabatier, Toulouse, France) who dis-

covered this highly interesting diapriid during his observations on myrmeco-

philic Diapriinae in France.

5. Plagiopria besucheti, n. sp. (Figs. 24, 25)

Female: Length 2.1 mm. Color dark chestnut brown with legs and anten-

nae yellowish.

Head dorsally (39:35) and laterally as in P. passerai, but pilosity on head

much sparser and hairs slightly longer; space between antennal shelf and

front orbits less longitudinally rugose; antenna (Fig. 24) with segments in

proportions: 4028: 1016; 1555. $7 Gi 7s 617s 6:7: 776i 556.53. 11.910 12:10;

—13.5:8; Al2 clearly narrower than A11; flagellum slightly more robust and

scape less reticulate than in P. passerai.

Mesosoma basically as in previous species; sides of pronotum in dorsal view

slightly more concave; only with two setae on pronotum close to scutal su-

ture; mesoscutum anteriorly conspicuously more bluntly rounded; scutellar pit

76 Contrib. Amer. Ent: Inst., vol. 20, 1983

very shallow, almost inconspicuous and virtually without median carina on

shield; propodeal flanges without wrinkles; legs more or less as in previous

species but mid tibia clearly more sinuate.

Petiole hardly longer than wide, slightly excavate anterodorsally, with much

shorter hairs laterally than in P. passerai and a fine longitudinal depression

laterally (Fig. 35); body of metasoma about 1.7 x as long as broad; T2 (85:57)

with only some few scattered long setae, all directed posteriorly (Fig. 35).

Male: Unknown.

Biology: Unknown.

Distribution: Turkey.

Type material: Holotype: Female, Turkey, Ordu, "Env. d'Ordu", May

17, 1967 (C. C. Besuchet) [MHN]. The following parts of the body are miss-

ing: fore and hind wings, apical trasomere of left fore tarsus and most of

left hind tarsus. Right antenna on slide.

6. Plagiopria huberi, n. sp. (Figs. 22, 36, 37)

Female: Length 2.0 mm. Color uniformly yellowish brown; wings hyaline.

Head in dorsal view globose (40:35) with prominent antennal shelf, rim-

like produced foreward and slightly concave medially; distinct longitudinal

ridge between antennal sockets; pilosity on temples sparse, indistinct; eyes

small, shorter than temples, glabrous, raspberry-like, with rather large omati-

dia; ocelli close together POL:LOL:00L = 5:5:12; frons slightly concave; ocellar

area somewhat elevated, with reticulation; head with scattered minute hairs;

in lateral view head triangular and space between orbits and apex of antennal

shelf with pronounced irregular reticulation extending down face; rest of head

shiny but with large-scaled, foveolate, shallow sculpture; eyes small, subcircu-

lar, eye height slightly greater than malar space; hypostomal carina lamel-

liform, pronounced; mouth small; palpi very short; antenna (Fig. 22) peculiar

in having distal funicular segments narrower than proximal ones and clava

inconspicuous; A2 - Al2 remarkably irregular in outline; antennal segments

in. proportions:.40:93.10-7.54 1010.32 0827 052 Bi 7p Os7: 1 B20.55 7262). 8263. 9575 9:75

12:6; scape and also pedicel with strong reticulation; A3 much shorter than

in previous two species and flagellum bristly.

Shoulders of prothorax more rounded, hardly angularly protruding; pronotum

with dense but small collar of ‘greyish pilosity and in‘ dorsal view with 5-6

hairs on each side near scutal suture as in Fig. 30; mesoscutum postero-

laterally (near tegular) and two anteromedial depressions shiny, rest of meso-

scutum with irregular, foveolate, longitudinal reticulation; mesoscutum sub-

medially on each side with irregular row of some short hairs; scutellar suture

distinct, rim-like; scutellar pit semicircular, wide, with rim-like anterior

margin and pit fan-like costate; scutellar shield parallel, finely margined

laterally, with distinct but delicate lamella-like edge posteriorly; scutellum

with irregular foveolate sculpture, less so on axillar parts and with delicate

median carina in posterior half of shield; blade-like flanges of propodeum

slightly shorter than in previous two species and median keel absent; sides

of mesosoma with same sculpture as on mesoscutum; fore wing as in

P. passerai but disc slightly denser pilose and apical cilia very short, only

Huggert & Masner: Diapriid wasps 77

about 1/14 of wing width; hind wing with cilia short, about 1/3 of wing width;

tarsi not particularly short (Fig. 36), distinctly compressed; femora and tibiae

compressed, femora with blade-like edges in basal half; tibiae much narrowed

basally compared to the other two species, club-shaped, gradually incrassate

towards apex, with delicate ventral and dorsal edges medially; row of comb-

like bristles on hind tibia less pronounced and mid tibia somewhat club-like

sinuate.

Petiole in dorsal view slightly wider than long (14:20), with rather bulging

sides, slightly narrowed anteriorly and here with fine margin; petiole lateral-

ly with rather distinct longitudinal groove on each side and with fine pilosity

all over, longer at sides (Fig. 37); body of metasoma 1.5 x as long as wide;

T2 (67:58) with scattered short hairs; S2 with broad median tongue bent

downwards anteriorly, bordered by short grooves.

Male: Unknown.

Biology: Unknown.

Distribution: Iran.

Type material: Holotype: Female, Iran, Tehran Prov., 8 km W Konchor,

July, 1978 (J. T. Huber) [CNC No. 17656] (left antenna on slide).

Remarks: Plagiopriahuberi differs distinctly from the other two species

in the structure of the antenna, legs and general sculpturing of the head and

mesosoma. This species exhibits several features paralleling character states

encountered among numerous New World ecitophilous diapriines, viz. body

usually light colored (xanthic or rufous), more or less sculptured, tarsi dis-

tinctly compressed laterally and antenna without distinct clava.

Etymology: This new species is named in honor of Mr. John T. Huber

(University of California, Riverside), who collected this wasp in Iran.

7. Trichopria myrmicae, n. sp. (Figs. 39-49)

Female: Length 2.4 mm. Black; legs entirely rufous, mandibles and

antennae rufous, with tips of mandibles and clava slightly darker; wings

hyaline. |

Head in dorsal view globose; antennal shelf only slightly protruding; tem-

ples rounded, subequal to length of eye; POL:LOL:00L = 11:6:18; head in later-

al view with eyes small, oval (15:23), about 2.5 x longer than malar space;

genae and temples with think greyish pilosity; hypostomal carina prominent;

palpal formula 5-2 (Fig. 40); head in frontal view subcircular with very wide

mouth, almost as wide as head; genae very short; mandibles unusually large,

unequal, with strong lower tooth and truncate upper one; right mandible with

upper tooth subbidentate (Fig. 43); clypeus with pronounced, deep transverse

cleft right above lower margin; face below antennal shelf with long bristles

in setigerous punctures, head otherwise with remarkably short hairs; antenna

_ (Fig. 39) notably short compared to body, hardly reaching hind margin of

propodeum; flagellar segments short, clava 3-segmented; Al to Al2 in propor-

tions: 34:8; 935.8; 935.53, G:6.3; 6:58: ..6:3.8: (6:0; 16,5762 6:55:62: 10:9: 10:9, 5: 14:9:

flagellar pilosity rather dense, shorter than width of segments.

78 Contrib. Amer. Ent. Inst., vol. 20, 1983

Prothorax with thick collar of greyish pilosity anteriorly (Fig. 44); setae

on mesoscutum unusually short, two placed in front of tegula and two, very

close to each other, placed about in middle on each half of mesoscutum;

axillae with group of 4-5 short hairs; scutellar pit transverse, lunate, with

two deeper pits laterally and these pits with some striae; scutellar shield

strongly convergent posteriorly, moderately arched with a strong median

carina in posterior 2/3; sides of scutellum with strong rim; median keel of

metanotum much elevated (Figs. 45, 46); propodeal plicae distinct and median

keel anteriorly elevated; propodeum and metapleura covered with dense long

whitish hairs; sides of pronotum below tegula with group of 5-6 short hairs;

mesopleura in lower quarter with arc of 6 erect hairs; fore wing with about

20 short setae on submarginalis + marginalis and wing proximally with rather

dense hairs (Fig. 41); longest marginal cilia about 1/8 of wing width; hind wing

with cilia almost half of wing width; legs unusual, femora strongly incrassate

with sharp edge ventrally; posterodorsal apex of fore tibia with remarkably

long, strong, curved spine (Fig. 47); inner margin of hind tibia with very well

developed row of dense, stiff bristles.

Petiole slightly longer than wide, completely covered by long greyish hairs;

body of metasoma (45:25) and T2 (33:25); T6 triangular and apex of metasoma

sharply pointed, due to projection of ultimate tip of last sternite; T3-T5

medially with transverse band of dense micropunctures (160 x), which under

magnification of 1600 - 4000x appear as rows of pegs (Figs. 48, 49).

Male: Unknown.

Distribution: USA (New York, St. Lawrence River Valley).

Biology: This is the first known diapriid associate of ants of the genus

Myrmica. All individuals were collected under stones in nests of Myrmica ?

americana Weber (det. L. Huggert) situated on open, south-facing, grassy

slopes on limestone. Only one or occasionally two individuals of T. myrmicae

were found per nest. Wasps were found only in about 20% of the examined

nests and they were seen moving slowly amidst the ants. However, the ants

did not seem to pay any particular attention to the wasps. One wasp was

brought into the laboratory for further observation in a formicarium. Again,

it was ignored by the ants. Similarly, we failed to observe any case of pet-

ting or trophalaxis but also no hostility towards the guest. We found it in-

teresting that the individuals of T. myrmicae occur in nests with grown larvae

(? L2) of the host ant. This also agrees with our experience with

Auxopaedeutes spp. associated with Solenopsis ants. Prior to the appearance

of larvae of Solenopsis, no individual of Auxopaedeutes wasps was observed

in the nests.

Variability: Very little variation has been noticed among the 12 individ-

uals except for the body size that varies between 2.3 - 2.6 mm.

Type material: Holotype: Female, U.S.A. (New York), Alexandria Bay,

April 25, 1978 (L. Huggert), palpi on slide [CNC no. 15595]. Paratypes:

4 F, same data as holotype and also of April 23, | F gold-coated, with an=

tenna and wings on slide [CNC]; 1 F, same data as holotype [USNM]; 6 F,

with same data as holotype and of April 23 [Hugg].

Huggert & Masner: Diapriid wasps 79

Remarks: In Kieffer's (1916) classification, T.myrmicaeyn. sp. would fall

into Ashmeadopria Kieffer, which more recently has been considered a

synonym of Trichopria Ashmead (Sundholm 1960). However, this new species

appears quite peculiar among all Holarctic species that we classify in the cur-

rent concept of Trichopria. The following combination of these peculiar

character states is considered important for the recognition of the new

species: remarkably short antennae compared to the large size of the body;

unusually short setae and their configuration on the vertex and mesothorax;

very wide mouth, broad clypeus and consequently long and powerful mandibles;

all the femora strongly incrassate and sharply edged ventrally; long, curved

spine on the dorsal apex of the fore tibiae; the row of bristles along the in- .

ner side of the hind tibiae unusually pronounced.

Since the present taxonomy of the genus Trichopria s. 1. in North

America is not satisfactory, it is virtually impossible to classify this new

species more precisely. However, T. myrmicae differs remarkably in the

character states mentioned above from all species of Trichopria in North

America, Europe and Africa, the types of which we have studied.

ACKNOWLEDGMENTS

Our thanks are due vo Dr. C. Besuchet (Geneve), Dr. G. Osella (Verona),

Dr. L. Passera (Toulouse) and Dr. F. Werner (Tucson) for kindly supplying us

with specimens. Dr. J. M. Campbell and Mr. M. J. Sharkey (Biosystematics

Research Institute, Agriculture Canada, Ottawa) reviewed the manuscript.

REFERENCES

Borgmeier, T. 1959. Sobre alguns Diapriideos myrmecophilos, principalmente

do Brasil (Hym. Diapriidae). Revta Ent., Rio de J. 10: 530-545.

Brown, W. L. In: B. J. Meggers etal. 1973. A comparison of the Hylean

and Congo-West African rain forest ant faunas. In: Tropical forest

ecosystems in Africa and South America: A comparative review.

Smithsonian Institution Press, Washington, D.C., pp. 161-185.

Brues, C. T. 1916. A new species of Lepidopria from North America. Psyche

23: 126-127.

Dessart, P. 1975. Materiel typique des microhymenoptera myrmecophiles de

la collection Wasmann depose au Museum Wasmannianum a Maastricht

Pays-Bas). Publicaties van het Natuurhistorisch Genootschap in Limburg

24: 1-99.

Donisthorpe, H.S.J. 1927. The guests of British ants, their habits and life

histories. George Routlege & Sons, Ltd., London. XXIII, 244 pp.

-Ferriere, C. 1929. Nouveaux Diapriides du Bresil, hotes de Eciton. Zool.

Anz. 82: 156<17k

Kieffer, J. J. 1916. Diapriidae. Das Teirreich. Vol. 44. Berlin, 627 pp.

Kozlov, M. A. 1966. New species of proctotrupids (Hymenoptera,

Proctotrupoidea) from the USSR. Trudy Zool. Inst. 37: 137-147 (in

Russian).

80 Contrib. Amer. Ent. Inst., vol. 20, 1983

Loiacono, M. S. 198]. Diapridos ecitofilos sinfilos de la region neotropical

(Hymenoptera - Proctotrupoidea - Diapriidae). Rev. Soc. Ent.

Argentina 40: 297-310.

Masner, L. 1964. Proctotrupoidea. In J. Jasic etal. : Hyphantria cunea,

Editorial Board Slovac Academy of Sciences; pp. 248-249.

Masner, L. 1977. A new genus of ecitophilous diapriid wasps from Arizona

(Hymenoptera: Proctotrupoidea: Diapriidae). Can. Ent. 109: 33-36.

Muesebeck, C.F.W. In K. V. Krombein et al. 1979. Catalog of Hymenop-

tera of America north of Mexico. Vol. 1, 1198 pp. Smithsonian In-

stitution Press, Washington, D.C.

Muesebeck, C.F.W. and Walkley, L. M. 1956. Type species of the genera

and subgenera of parasitic wasps comprising the superfamily

Proctotrupoidea (Order Hymenoptera). Proc. U.S. Nat. Mus. Washington

105: 319-419.

Pschorn-Walcher, H. 1957. Zur Kenntnis der Diapriinae (Proctotrupoidea) der

Wasmann-Sammlung. Mitt. Schweiz. Ent. Gesell. 30: 73-88.

Sundholm, A. 1960. On Diapria Latreille and allied genera (Hym., Diapriidae)

Opuse. Ent, 25: 213-225.

Szeleny, G. 1957. Some new data on the hymenopterous parasites of

Hyphantria cunea Drury. Ann. Inst. Prot. Plant. Hung. 7: 295-312.

Wilson, E. O. 1972. The Insect Societies. Belknap Press, Cambridge, Mass.

I-X, 548 pp.

Wing, M. W. 1951. A new genus and species of myrmecophilous Diapriidae

with taxonomic and biological notes on related forms (Hymenoptera).

Trans. R. Ent. Soc. Lond. 102: 195-210.

Huggert & Masner: Diapriid wasps

©)

AL P

ager he TASe

ALECTOMY BEGINS

EMERGENCE

ALATED ADULTS

PL.

22 (S ALECTOMY COMPL

For

)

OVIPOSITION

Fig. 1. General scheme of the life cycle of symphilic

diapriid wasps associated with ecitonine ants.

Fig. 1. General scheme of the life cycle of symphilic

diapriid wasps associated with ecitonine ants.

Pigs, 20.

mellea Ashmead. 3,

imitatrix Wasmann.

pedestris HKieffer.

passerai, n. sp.

Contrib.-Amer. Ent. Insts) vol. 20,

Petiolar region,

5 ESD

5 Y LU Ei

2 Waar eK,

SEE Zs

te Ce. é

Ws Oly

‘ es!

3 CY >,

Vs :

Scat

Niet

“ia,

Bruesopria sp.

Auxopaedeutes sp.

females. 2, Myrmecopria

Ecitovagus gibbus Masner.

1983

4, Solenopsia

Lepidopria

8, Plagiopria

FAnS, Geile «

(Schemeaticogs.+.8%,

dopria pedestris

Auxopaedeutes sp.

Huggert & Masner: Diapriid wasps

Lateral view of female petiolar region

Solenopsia imitatris Wasmann.

Kieffer. 11, Bruesopria sp.

12;

Lepi-

84 Contrib. Amer. Ent. Inst., vol. 20, 1983

Figs. 13-193 Leabidopria noctivaga, mn. sp. F. and

Townesella marjoriae, n. gen. et sp., F. 13, L. noctivaga,

lateral view.” 14, °T. “‘marjoriage, “lateral view. 15, Ll.

noctivaga, dorsal view. 16. T. marjoriae, dorsal view.

17 Ly. tectigava, petiolar region’ (Caoblique view). To,’ ts

mar joriae, potiolar region (dorsal view). 19, T. marjoriae,

temple and pronotal shoulder (oblique view).

Huggert & Masner: Diapriid wasps 85

Figs. 20-24. Female antennae: 20, Townesella marjjoriae.

21, Labidopria noctivaga. 22, Plagiopria huberi, n. gen. et

sp. 23, P. passerai, n. sp. 24, P. besucheti, n. sp.

86 Contrib. Amer. Ent. Inst., vol. 20, 1983

Figs. 25-29. Cordylocras mirabilis Kozlov, F. and

Ecitovagus gibbus Masner, M.: 25, C. mirabilis, head lateral

view. 26, GCG. mirabilis, head dorsal. wiew. </, C. mirabilis,

pronotum and mesoscutum, dorsal view (after Kozlov 1966,

modified). 28, E. gibbus, antenna. 29, E. gibbus, A5-A5.

Huggert & Masner: Diapriid wasps

. Figs.

Fy bodys

antenna.

P. huberi

30-37.

FAs Be

hie. Md

ae ee

Plagiopria,

head lateral view.

CO OT SS"

inte wa

Tl.

gen.

MEF

Be Liles

besucheti,

P. huberi, F.

30,

passerai,

hind degac 33557 Ms

F. metasoma. 36,

metasoma.

88 Contrib. Amer. Ent. Inst., vol. 20, 1983

Figs. 38-41. Plagiopria passerai, n. gen. et sp. F.

and Trichopria myrmicae, n. sp. F.: 39, P. passerai, base

of fore wing. 39, T. myrmicae, antenna. 40, T. myrmicae,

padpi..49)° bo myppicae, base of fore. wing.

Huggert & Masner: Diapriid wasps 89

Figs. 42-49. Trichopria myrmicae, n. sp. F.(SEM coated

specimen): ’ 42) Heed C106" «0. 63. Mouth 1256 40% °- 4s, Mesos

soma dorgal view C132 x). 45, Propodeum, posterior view ©€157x).

46, Scutellum-propodeum, lateral view (224 x). 47, Fore

leo’ (233 x)eic Be yoboroecul pture, epek ot f2 0 166: wc

Microsculpture,apex of T2 (4040 x).

A NEW GENUS OF TETRASTICHINAE FROM THE NEW WORLD

(HYMENOPTERA: CHALCIDOIDEA: EULOPHIDAE)

C. M. Yoshimoto

Canadian Forestry Service

Department of Environment, Ottawa

Abstract

A new genus and species of Tetrastichinae, Henryana magnifica,

related to the genus Tetrastichus Haliday (Chalcidoidea: Eulophidae), is

described from specimens from Nova Teutonia, Brazil.

Specimens representing a new species and genus were found among the

many chalcid wasps collected by Fritz Plaumann and housed in the Canadian

National Collection, Ottawa. The elongate head, thorax and gaster, long ovi-

positor and lanceolate fore wing separate members of this taxon from other

chalcidoids. |

The new genus belongs to the family Eulophidae, subfamily

Tetrastichinae and is related to the genus Tetrastichus Haliday. Recent

publications on this subfamily (Boucek 1977, De Santis 1979, and Burks 1979)

were reviewed but none dealt with this genus.

HENRYANA, new genus

Head elongate in frontal view, about 3 times as long as wide, longer

than mesoscutum; face narrowing posteriorly; clypeus minute, narrow and long,

its anterior margin rounded; antenna inserted at level of ventral margin of

eyes; antennal scape linear, reaching frontal ocellus; club as long as 3 funicular

segments, anelli tiny, 2-segmented; eye ovate, bare; ocelli in form of equi-

lateral triangle; malar space about half as long as length of eye (Fig. 1).

Neck elongate, about 3 times as long as wide, membraneous medially;

pronotum elongate, subrectangular in dorsal view, dorsal area non-sclerotized,

its margin keeled, pleural area non-sclerotized; mesoscutum elongate, 3.5

times as long as wide, tapering anteriorly, its lateral lobe about 1/3 length

of mesoscutum, not reaching pronotum (Fig. 2); notauli broadly grooved, about

1/3 as long as mesoscutum; scutellum short, about, 3.5 times as long as wide,

about 1/3 as long as mesoscutum; axilla subtriangular, slightly in front of

tegula; prepectus membraneous, forming a large subtriangular area ventrally;

dorsellum large, triangular; propodeum with y-shaped membraneous furrow, fol-

lowed by bell-shape depression continuing to base of petiolar insertion;

mesepicnemium narrow and long; mesepisternal plate large, subrectangular.

Fore coxa elongate, cone-shaped, nearly as long as prosternal plate; mid and

hind basitarsus equal in length, mid tarsal spine minute, barely visible.

*Seaanded to Biosystematics Research Institute, Agriculture Canada, Ottawa

KIA OC6.

Yoshimoto: Tetrastichinae 91

Fore wing lanceolate, about 8 times as long as wide, extending to ter-

gite 6 (Fig. 1); marginal vein about 3 times as long as submarginal vein;

stigmal vein about 1/3 as long as submarginal vein; marginal cilia longer than

widest part of fore wing. Hind wing blade-like, with 3 hamuli.

Petiole sessile, narrow, crescentic (Fig. 2). Female gaster 3.5 times as

long as wide, slightly shorter than thorax; male gaster as long as thorax;

terga 1-5 with deep, median longitudinal groove; posterior margin of tergum

1, with deep u-shaped emargination; lengths of terga 1-7 in ratio 0.5: 0.5: 2:

4: 3: 2: 1. Ovipositor long, about as long as terga 2 and 3 combined.

Type-species: Henrya magnifica,n. sp., present designation.

I am delighted to name this genus in honor of Dr. Henry Townes.

Females of this genus are similar to Tetrastichus Haliday but differ in

having the clypeus vertically narrow, subrectangular, its anterior margin round—

ed; eyes bare; antennal scape of female linear, reaching the frontal ocellus;

mesoscutum elongate, its lateral lobe about 1/3 the length of mesoscutum, not

reaching the pronotum; dorsellum large, triangular; fore wirg lanceolate, 8

times as long as wide, the marginal vein 3 times as long as the submarginal

vein; dorsal surface of pronotum and entire prepectus membraneous; propodeum

with y-shaped membraneous furrow dorsomedially; petiole sessile, narrow,

crescentic; basal margin of tergum | forming a deep u-shaped emargination,

terga 1-5 deeply grooved medially and ovipositor as long as terga 2 and 3 com-

bined.

Henryana magnifica, n. sp. (Figs. 1, 2)

Female: Length 2.0 mm. Brown to dark brown; head yellowish brown,

except frontal crest and scrobe brownish; antennal scape yellowish to yellowish

brown, dorsal margin darker, pedicel and flagellum dark brown; propleuron

yellowish brown, mesepisternum yellowish brown, tarsi slightly darker.

Antennal scape smooth, linear, 7 to 8 times as long as wide; ventral mar-

gin with | or 2 minute setae; lengths of antennal segments excluding annellus

in ratio 3: 1: 1: 0.9: 1: 0.9: 0.7 (Fig. 1); malar space smooth, longer than wide;

malar groove present; vertex and frons forming membraneous area, frontal and

interantennal crest forming acute triangular sclerotized plate, its anterior part

reaching middle of frons, scrobe forming elongate u-shaped area.

. Mesoscutum coriaceous with median line (Fig. 2); scutellum with single

seta mid-distance from anterior margin; dorsellum large, triangular, posterior

margin extending into propodeal area; propodeum smooth, spiracle tiny, touch-

ing anterior margin, callus with single seta, postero-lateral area with single

seta, petiolar insertion u-shaped; median carina absent, thin plica reaching

spiracle.

Gaster coriaceous, lateral areas of tergum with scattered setae; pygostyle

with two short and single long bristles.

Male: Unknown.

Type: Holotype, female, Nova Teutonia, Brazil, 27°11'S, 52°23'W,

X-XI1.1972, Fritz Plaumann. (CNC No. 17719). Paratypes: 4 females, same

data as holotype.

92 Contrib. Amer. Ent. Inst., vol. 20, 1983

ACKNOWLEDGEMENTS

My thanks to my colleagues in the Biosystematics Research Institute,

A. Smetana, V. Behan-Pelletier and M. Sharkey for their comments and Susan

Rigby for the illustrations.

REFERENCES

Boucek, Z. 1977. Descriptions of Tachinobia gen. n. and three new species

of Tetrastichinae (Hymenoptera: Eulophidae), with a tentative key to

genera. Bull. Ent. Res. 67: 17-30.

Burks, B. 1979. In Krombein, et al. Catalog of Hymenoptera in America,

north of Mexico. Smithsonian Press. 1: 967-1022.

De Santis, L. 1979. Catalogo de los himenopteros calcidoideos de America

al Sur de los Estado Unidos. Pub. Especial, Provincia de Buenos Aires,

Comision de Investigaciones Cientificas. La Plata. 488 pp.

Yoshimoto: Tetrastichinae

Figs. 1-2. Henryana magnifica, nm. sp.-: WV

habitus Lateral ‘view.

Dorgal’ view ofvEnorax.

Female,

MARJORIELLA, A NEW NEOTROPICAL GENUS OF

AGATHIDINAE (BRACONIDAE, HYMENOPTERA)

M. J. Sharkey

Biosystematics Research Institute

Agriculture Canada, Ottawa K1A 0C6

Abstract

Marjoriella, a new genus of Agathidinae from the Neotropical region,

is described. Four species, all new are included: M. ancha, M. beni,

M.henryi and M.townesorum. A cladistic analysis is presented suggesting

the clade, Megagathis Kriechbaumer plus Labagathis Enderlein, as the sister

group of Marjoriella.

INTRODUCTION

Little work on the Neotropical fauna of the braconid subfamily

Agathidinae has been done in the last 60 years. Brullé (1846), Enderlein

(1918), Szépligeti (1904), and Viereck (1912a, 1912b, 1913), described several

Neotropical agathidinae genera; however, the characters of none of their

genera agree with those of the new genus described here. Intergeneric re-

lationships are based on cladistic methods (see Hennig 1966 and Wiley 1981).

Character states have been polarized using outgroup analysis (see Waltrous

and Wheeler 1981).

Abbreviations

The following abbreviations are used for institutions in which type-

material of the new species described in this paper are deposited.

A.M.N.H.: American Museum of Natural History, New York, New York.

C.N.C.: Canadian National Collection, Ottawa, Ontario.

C.P.H.: Center for Parasitic Hymenoptera, Gainesville, Florida.

Wing venation: (Fig. 3e): The Comstock-Ross system of nomenclature

for wing venation has been used.

The venation of Marjoriella is not fully understood in the area of the

areolet. It is not clear whether the vein labelled RS; (Fig. 3e) is, in fact,

the 2nd radial sector or simply an adventitious vein. This confusion will be

resolved only after a cladistic analysis of the entire subfamily allows the

origin of the vein to be elucidated. If the vein is part of the ground plan

for the Agathidinae, then there is support for my hypothesis that it repre-

sents a remnant of the second radial sector. Conversely, a cladistic analysis

may show the vein to be a derived adventitious vein not homologous with

‘'RS2.

MARJORIELLA, new genus

Type-species: Marjoriella henryi, n. sp.

Gender of the genus: Feminine.

Length of body about 10.0 mm. Head: Head wider than long (Fig. la);

face moderately long but not rostriform; malar space shorter than eye height;

maxillary palpus 5-segmented; labial palpus 4-segmented, segments subequal

Sharkey: Agathidinae 95

in length; frontal pit deep, with frontal and marginal carinae; marginal carinae

strongly elevated, between and laterad of antennal sockets; vertex (in dorsal

view) not excavated; occiput with strong lateral carinae; gena very wide, about

equal to eye width, with sharp groove posteroventrally for reception of lateral

portion of pronotum; gena forming acute angle at its apex; antenna long, equal

to or longer than body, 50-to 54-segmented.

-Mesosoma: Pronotum with large anterior flange occupying space between

carinae of occiput (Fig. 3a); pronotum strongly compressed posterodorsally an-

terior to mesoscutum; sternalus strongly foveolate; mesoscutum smooth, notauli

absent; scutellum smooth, without carinae; posterior portion of mesoscutum,

scutellum and metanotum on same plane; metanotum smooth medially, without

sculpture or armature; propodeum (Fig. 3c) flat; with deep groove antero-

medially and small depression centrally, otherwise smooth and glabrous medial-

ly; propodeal spiracles large and oval; propleura with pair of projections extend-

ing between middle coxae (Fig. 3b); hind coxal cavities open; basal tooth of

tarsal claws forming sharp angle (Fig. 3d); spines absent from outer surface

of middle tibia; areolet of fore wing sessile (Fig. 3e), 4 sided, with small 2nd

radial sector; 2rm of hind wing not indicated.

Metasoma: All terga smooth, without sculpture except for weak V-

shaped groove sometimes present on tergum 2; tergum | wide, length about

equal to apical width; ovipositor very short, decurved.

Diagnosis: These are large agathidines readily separated from others

by the following combination of characters: Occiput with strong lateral

carinae; areolet sessile and 4-sided with a small segment of 2nd radial sector

present, propleura and mesopleura with projections ventrally between the fore

and middle coxae respectively; propodeum flat, not convex, with a deep longi-

tudinal anteromedial groove.

Distribution: Restricted to South America; see map (Fig. 2).

Etymology: I am pleased to name this genus after Dr. Marjorie Townes

who has worked at Henry's side for many years and contributed greatly to his

work.

Relationships: Cladistic methods were used in this study to hypothe-

size the phylogenetic relationships of Marjoriella. Character states were

polarized using outgroup comparison, the outgroup being the remaining taxa of

Agathidinae.

Marjoriella is highly autapomorphic in many characters, such as the

shape of the areolet and the sculpture of the propodeum, that are useful in

making suprageneric groupings in another genera within the Agathidinae. In

spite of this there are two convincing synapomorphies uniting Marjoriella

with Megagathis Kriechbaumer and Labagathis Enderlein. Members of both

of these genera, like Marjoriella, have large anterior flanges on their pronota.

The second synapomorphy is the strong lateral carinae of the occiput. These

carinae are not homologous with occipital carinae. In Megagathis these

carinae are reduced and rounded. No other braconids have anterior flanges

on the pronotum or lateral carinae on the occiput. The phylogram (Fig. 1)

illustrates the relationships of these three genera. Megagathis plus

Labogathis is the sister group of Marjoriella.

96 Contrib. Amer. Ent. Inst., vol. 20, 1983

The monophyly of Marjoriella is supported by the following synapo-

morphies of its component species:

1. Spines on pro- and mes-episternum present (Fig. 3b).

2. Unique shape of areolet (Fig. 3e).

3. Unique shape of propodeum (Fig. 3c).

Biology: All agathidines, whose hosts are known, parasitize lepidopteran

larvae, and Marjoriella is likely no exception, though there are no host

records for this genus. Because of the short size of their ovipositors they may

parasitize exposed larvae, rather than those concealed in leaf rolls, thick webs,

or in stems. With this in mind and due to their rather large size | hypothe-

size their hosts to be noctuids and probably members of the subfamily

Catocalinae. The Catocalinae are the dominant group of Lepidoptera in Neo-

tropical rainforests, comprising the vast majority of the 16,000 described

species of noctuids from the Neotropical region.

KEY TO SPECIES OF MARJORIELLA (Females)

LS SWweies Banded Geo Wis EE ke ee Pe Cea St cae wut v4

Wings not banded, yellowish with darker stigmal area ..........-..

4. townesorum, n. Sp.

Ces” SUMS, cue SRNR Soabey Gm het YeMRO Veen’ | Aika SMe ee ie haut A, Oe ae ENO RS Yt ER. Ae SAT, Se om Jot kek UMM SiR Tend

2. Propodeum black, with areolae mesad of spiracles ... .2. beni, n. sp.

Propodeum testaceous, smooth between spiracles except for medial

RSVR ES. i SES SS CU I Us a CR See as PK 3

3. Terga 2+3 wider than long, length/width = 0.8; hind femur incrassate,

ET ON Sr ee ANS LR ate et TIE Bla wot: aha. One 8D,

Terga 2+3 longer than wide, length/width = 1.4; hind femur of normal

PTOMOLE IONE. 1OMeUNWIKTR Bie i sk TRA See i CO a OREO FL ae.

1. Marjoriella ancha, n. sp.

The adult female of this species is most similar to that of M. beni but can

easily be distinguished by its lighter coloration and the lack of areolae in the

area between the propodeal spiracles.

Female: Length 10.0 mm. Black and testaceous; head including antennae

black; anterior portion of mesosoma black; metapleuron brown; propodeum

testaceous, all coxae black laterad, testaceous mesad, legs black, brown and

testaceous; metasoma testaceous except black apically.

Antenna 54 segmented. Metapleuron entirely areolate rugose; hind femur

incrassate, length/width = 2.5; fore wing banded yellow and black, black bands

positioned apically and in stigmal area; 1 RS of normal proportions | RS / 1

RS 9 = 1.6. Metasomal terga rather wide; tergum | wider than long, length /

apical width = 0.9; terga 2+3 wider than long, length / width = 0.8; ovipositor

short, ovipositor sheath 1/5 length of hind femur.

Male: Unknown.

Holotype: F, Surinam, Paramaribo, 13.1X.1963, D.C. Geijskes (C.P.H.).

Sharkey: Agathidinae o7

Etymology: Ancha means wide in Spanish and refers to the wide body di-

mensions of female of this species.

2. Marjoriella beni, n. sp.

The adult female of M. beni is similar to that of M. ancha in general size

but is easily distinguished from the latter species by darker coloration of and

presence of areolae between the propodeal spiracles.

Female: Length 10.0 mm. Black except fore tarsi, parts of middle tarsi

and hind trochanters testaceous; hind legs medially and metasoma anteriorly

reddish. |

Antennae 52-segmented. Metapleuron entirely areolate rugose, hind femur

incrassate, length/width = 2.8; fore wing mostly infuscate, yellowish basally

and with hyaline band distad of stigma; 1 RS rather short, | RS / IRS» 212.3;

propodeum with oval depressions mesad of and subequal to propodeal spiracles.

Metasomal terga wide; tergum | wider than long, length / apical width = 0.77;

terga 2+3 about as wide as long, length / width = 0.95; tergum 2 with a strong

transverse V-shaped groove, glabrous anterior to groove, setose posteriorly;

Ovipositor sheath short, about 1/5 length of hind femur.

Male: Unknown.

Hologype: F, Bolivia, Beni, Rio Itenez, opposite Costa Marques (Brazil),

4-6.XII1.1964, Bouseman + Lussenhop (AMNH).

3. Marjoriella henryi, n. sp.

The adult female of M. henyri is most similar to M. townesorum but is dis-

tinguished from the latter by presence of a dark band in the stigmal area of

the fore wing.

Female: Length 9.5 mm. Testaceous except: head and antennae black,

propleuron black, pleural regions of thorax brown to black, all coxae black

laterad, testaceous to yellow mesad, most of middle tibia and hind tarsi brown,

propodeum brown, metasoma black apically.

Antennae 5l-segmented. Metapleuron partly smooth but with some large

rugose areolae; hind femur not too wide, length / width = 3.2; fore wing banded

yellow and black, black bands positioned apically and in stigmal area; 1 RS rather

long, 1 RS /1RS = 2.2. Metasomal terga of normal dimensions; tergum |

as wide as long, terga 2+3 longer than wide, length / width = 1.4; ovipositor

short, ovipositor sheaths about 1/5 length of hind femur.

Male: Unknown.

Variation: The paratype F differs from the holotype only in its lighter

coloration i.e. mesopleurae and propodeum testaceous.

Holotype: F, Brazil, Mato Grosso, Sinop, X.1974, Malaise trap, M. Alvarenga.

Paratype: F, same data as holotype (CNC).

Etymology: ‘This species is named for Dr. Henry Townes, in honor of his

distinguished career in entomology.

98 Contrib. Aner.” Ent: Inst.;°voli 20, 1983

4. Marjoriella townesorum, n. sp.

The adult female of M. townesorum is most similar to that of M. henryi,

but may easily be distinguished from it by the narrower metasoma and the

lack of a dark band in the stigmal area of the fore wing.

Female: Length 10.0 mm. Testaceous except head including antennae

black, pronotum black anteriorly, hind tarsi brown, metasoms black distally.

Antennal number ? (broken). Metapleuron mostly smooth with weak rugae

peripherally; hind femur of normal proportions, length / width = 3.4; fore wing

darkening apically but without dark median band as in (Fig. 3e); | RS of nor-

mal proportions, | RS / 1 RS»5= 1.7. All metasomal terga smooth and shining,

without grooves; tergum | long, longer than wide, length / apical width = 1.1;

terga 2+3 long, longer than wide, length / width = 1.2; ovipositor very short,

not extruded; ovipositor sheaths barely visible in holotype, small, less than 1/5

length of hind femur.

Male: Unknown.

Holotype: F, Brazil, Mato Grosso, Sinop, 12°31'S, 55°37'W, X.1974,

M. Alvarenga (CPH).

Etymology: I name this species in honor of Henry and Marjorie Townes

and their union.

ACKNOWLEDGMENTS

I thank Dr. W. R. Mason for helpful comments and Dr. Lubomir Masner

and Dr. Don Lafontaine for reviewing the manuscript (all from the Biosystem-

atics Research Institute, Ottawa).

REFERENCES

Brulle, A. 1846. In: Lepeletier De St. Fargeau, A.L.M. Suites a Buffon:

Histoire naturelle des insectes: Hymenopteres. Paris. Roret, 4, 689 pp.

Enderlein, G. 1918 (1920). Zur Kenntnis aussereuropaischer Braconiden. Arch.

Naturgesch. 84(A) (11): 51-224.

Hennig, W. 1966. Phylogenetic Systematics. University of Illinois Press,

Urbana.

Szepligeti, G. V. 1904. Hymenoptera, Fam. Braconidae (Premiere partie)

Genera Insectorum, 22 and 23.

Viereck, M. L. 1912a. New genus and species of Hymenoptera of the family

Braconidae from Panama. Smithsonian Misc. Coll. 59(5): 1-2.

Viereck, M. L. 1912b. Contributions to our knowledge of bees and ichneumon-

flies, including the description of twenty-one new genera and fifty-seven

new species of ichneumon-flies. Proc. U.S. Natl. Mus. 42: 613-648.

Viereck, M. L. 1913. Descriptions of twenty-three new genera and thirty-one

new species of ichneumon-flies. Proc. U.S. Nat. Mus. 46: 359-386.

Watrous, L. E. and Wheeler, Q. D. 1981. The out-group comparison method

of character analysis. Syst. Zool. 30: 1-11.

Wiley, E. O. 1981. Phylogenetics, the theory and practice of phylogenetic

systematics. John Wiley and Sons. New York.

Sharkey: Agathidinae

Marjoriella

Megagathis

Labagathis

Fig. 1. Proposed phylogeny of Marjorie//a and its sister group Megagathis

plus Labagathis.

Fig. 2. Distribution map of Marjoriella spp.

Synapomorphies

Pronotum with a

large anterior flange

Occiput with lateral

carinae

Metasoma petiolate

Margin (lateral and

marginal carinae) of

frons lost.

M. ancha

M. beni

M. henrya

M. townesorum

100 Contrib. Amer. Ent. Inst., vol. 20, 1983

Fig.3

Fig. 3. A, Marjoriella ancha, posterodorsal view of

head and anterior mesosoma. APF, anterior pronotal flange;

FC, frontal carina; Mn, mesonotum; MC, marginal carinae;

OG; (ABT EPaL i Carine oF Aeetouts Pr, Seronotum,

B, M. townesorum, ventral aspect of mesosoma. C_,, coxa 1; Mp,

mesopleuro.; P Mp, projections of mesopleuron; Pp, propleuron;

P Poy projections of propaledron. .0, RM. eancha, propodeum.

D, M. henryi, tarsal claw of hind leg. E, M. beni, fore wing.

Ar, areolet; M, medial vein; r-m2, 2nd abscissa of radial-medial

cross vein; I RS, tet abetiage of radial sectiar; Ro), dst

vranch oF radial sectors Thae, et: aheoless of end @reanch: of

radial sector; 2RS2¢, 2nd abscissa of 2nd branch af radial sector.

REVISION OF SPECIES OF WESTERN PALEARCTIC ICHNEUMONIDAE

DESCRIBED BY FRENCH AUTHORS

Klaus Horstmann

Zoologisches Institut der Universitat, Wurzburg

Germany

Abstract

58 species of Western Palearctic Ichneumonidae described by Brullé,

Dufour and Perris, Boyer de Fonscolombe, Lucas, Giraud, Laboulbéne, and

Sichel are revised, 34 lectotypes are designated, and 25 new synonyms are in-

dicated.

This paper is the second result of my revisory work at the National Mu-

seum of Natural History in Paris (see Horstmann 1980). It is intended as a

prelude to a catalogue and reclassification of the Western Palearctic

Ichneumonidae.

ACKNOWLEDGMENTS

Mademoiselle Dr. S. Kelner-—Pillault and Madame M. Lachaise (Muséum

National d'Histoire Naturelle, Paris) helped me during my work at the Paris

Museum. Dr. J.-P. Aeschlimann (Commonwealth Scientific and Industrial Re-

search Organisation, Biological Control Unit, Montpellier), Dr. Paul Dessart

(Institut Royal des Sciences Naturelles Belgique, Bruxelles), E. Diller

(Zoologische Staatssammlung, Miinchen), Dr. M. Kak (Muzeum Przyrodnicze,

Wroclaw), Dr. P. Passerin d'Entreves (Istituto di Zoologia Sistematica dell'

Universita, Torino), and Dr. H. D. Volkart (Naturhistorisches Museum, Bern)

sent types or other specimens for comparison. Dozent Dr. M. Fischer (Natur-

historisches Museum, Wien) searched for types at the Vienna Museum.

E. Diller (Zoologische Staatssammlung, Miinchen), R. Hinz (Einbeck), Dr. G.

van Rossem (Plantenziektenkundige Dienst, Wageningen), and H. Schnee

(Markkleeberg) assisted with determinations of species. I am greatly indebted

to all of them.

BRULLE

In two publications, A. Brullé (1832, 1846) described 14 new species of

Palearctic Ichneumonidae. Brulle described most species from materal of the

National Museum of Natural History in Paris, which he rearranged, and which

is preserved in the Paris Museum as a separate collection near the name

"Brullé Collection". In the meantime, however, this collection was partly re-

arranged, and partly transferred to the general collection of Ichneumonidae,

and partly destroyed. Some of the new species were described from material

of the Serville Collection, which was bought by Spinola in 1847, and which is

now preserved in the Museum of Systematic Zoology of the University of Turin

(Italy), as a part of the Spinola Collection of Hymenoptera (Casolari and

Casolari Moreno 1980).

Pimpla longicauda Brulle

Pimpla longicauda Brullé, 1832: 379. Type(s) lost.

Identity: Exeristes roborator (Fabricius, 1793) (Oehlke, 1967: 1).

101

102 Contrib. Amer. Ent. Inst., vol. 20, 1983

Cryptus laticinctus Brullé

Cryptus laticinctus Brulle, 1832: 379. Type(s) lost.

Identity: Chaetomastrus laticinctus (Brulle 1832), syn. Hemiteles

hirticeps Thomson, 1885, syn. Hemiteles hirtus Brauns, 1888. New synonym.

Lectotype of Hemiteleshirtus Brauns hereby designated: F, "Brauns

det.", "2613", "VII. 87 Susten". Museum of Natural History, Bern.

Trachynotus humeralis Brullé

Trachynotus humeralis Brullé, 1832: 381. Type(s) lost.

Identity: Anomalon foliator (Fabricius,1798). New synonym.

Rhyssa marginalis Brulle

Rhyssa marginalis Brullé, 1846: 79. Type (? holotype): F, "Rhyssa

marginalis Br.". Paris Museum, general collection under Rhyssa persuasoria.

Origin unknown, but Townes (1961: 174) supposed that the type came from

France.

Identity: Rhyssa persuasoria (Linnaeus,1758) (Townes 1961: 174).

Pimpla bilineata Brullé

Pimpla bilineata Brullé, 1846: 98. Type(s) lost.

Identity: Pimpla spuria Gravenhorst, 1829. New synonym. Interpreted

in accordance with Lucas (1849: 323) and Tosquinet (1896: 284), whose material

came from the type locality (Algeria) and agrees with Brullé's original descrip-

tion.

Pimpla abdominalis Brullé

Pimpla abdominalis Brullé, 1846: 99. Primary homonym of Pimpla

abdominalis Gravenhorst, 1829. Type (? holotype): ? M, green round label,

"Museum Paris Egypte Bove 1833". "abdominalis Br." (only parts of thorax and

gaster preserved). Paris Museum, general collection under Tromatobia

variabilis.

Identity: Tromatobia variabilis (Holmgren,1856) (Shaumar 1966: 444).

Pimpla ephippium Brulle

Pimpla ephippium Brullé, 1846: 99. Type (? holotype): M, green round

label, "Museum Paris Egypte Bove 1933", "ephippium Br." (only parts of head,

thorax and gaster, one base of antenna, and the wings preserved). Paris

Museum, general collection under Itoplectis.

Identity: Itoplectis melanocephala (Gravenhorst, 1829) (Perkins 1941: 646).

Euceros dimidiatus Brullé

Euceros dimidiatus Brulle, 1846: 117. Lectotype hereby designated: M,

"M", Euceros dimidiatus Br." (head, front part of thorax, and hind part of

gaster lacking). Turin Museum, Spinola Collection. The other syntypes men-

tioned in the original description could not be found.

Identity: Euceros albitarsis Curtis, 1837 (Giraud 1857: 166).

Horstmann: Ichneumonidae 103

o e e ?

Metopius pinatorius Brulle

Metopius pinatorius (Bosc inlit.) Brullé, 1846: 120. Lectotype desig-

nated by Townes (1961: 171): M, "Museum Paris ex coll. Bosc", "TI.

pinatorius n. Bosc Carolina" (old label from Bosc collection). Paris Museum,

general collection, Nearctic material under Metopius. Townes and Townes

(1959: 99) supposed that the type came from Europe.

Identity: Metopius pinatorius Brulle, 1846 (Townes and Townes !959: 99).

Odontomerus striatus Brullé

Odontomerus striatus Brullé, 1846: 123. Lectotype hereby designated:

F, "Museum Paris", "Odontomerus striatus Br." (most parts of one antenna,

one front wing, and hind part of gaster lacking). Paris Museum, general col-

lection under Ischnoceros rusticus.

Identity: Ischnoceros rusticus (Geoffroy in Fourcroy 1785). New

synonym.

Ophion arcuatus Brulle

Ophion arcuatus Brullé, 1846: 146. Type (? holotype): "China", " Ophion

arcuatus Br.". Paris Museum, Brulle Collection. Townes (196l: 172) supposed

that the type came from France.

Identity: Enicospilus undulatus (Gravenhorst, 1829). (Townes 1961: 172).

Cryptus bovei Brulle

Cryptus bovei Brullé, 1846: 199. Lectotype hereby designated: F, green

round label, "Museum Paris Egypte Bove 1833" (one flagellum, some tarsi, and

underside and hind part of gaster lacking). Paris Museum, found in the Brulle

Collection, and transferred to the general collection under Cryptus.

Identity: Synechocryptus bovei (Brulle, 1846) (van Rossem 1969b: 371).

Ischnoceros maculipennis Brullé

Ischnoceros maculipennis Brullé, 1846: 261. Type(s) lost.

Identity: Chirotica decorator (Villers 1789) (Horstmann, in press).

Secondary homonym of Hemiteles maculipennis Gravenhorst, 1829.

-Ichneumon xanthomelas Brullé

Ichneumon xanthomelas Brulle, 1846: 309. Primary homonym of

Ichneumon xanthomelas Gmelin, 1790. Type (? holotype): F, "Museum Paris

Barbarie Guyon 253-39", blué round label, "Ichneumon xanthomelas Br.".

Paris Museum, general collection under Stenophorus instructor.

Identity: Ctenochares bicolorus (Linnaeus,1767) (Shaumar 1966: 449).

DUFOUR AND PERRIS

In a paper on the Hymenoptera living in dried bramble sticks, L. Dufour

and E. Perris (1840) described six new species of French Ichneumonidae. The

Dufour Collection is preserved in the Paris Museum, but some specimens from

that collection had been transferred to the Giraud Collection and to the general

104 Contrib. Amer. Ent. Inst., vol. 20, 1983

collection of Ichneumonidae of the Paris Museum, and some others had been

acquired by Spinola and are now found in the Spinola Collection in the Turin

Museum. The Perris Collection is preserved in the Zoological Insitute of the

National High School of Agriculture in Montpellier. I did not see it, but

Aeschlimann sent me material of some species. Usually both authors used la-

bels only with catalogue numbers, and only labelled the first specimen of a

series. Aeschlimann sent me a list of notes referring to those numbers from

the catalogue of the Perris Collection, but I do not know the meaning of the

numbers used by Dufour.

Ichneumon gyrator Dufour et Perris

Ichneumon gyrator Dufour et Perris, 1840: 40. Primary homonym of

Ichneumon gyrator Thunberg, 1822. Lectotype labelled by Sawoniewicz and

hereby designated: F, without original label. Turin Museum, Spinola Collec-

tion under the label "Cryptusgyrator Duf. F. - n. sp. ? D. Dufour S. Sever.".

Some other specimens, which may be syntypes, are in the Dufour Collection

in Paris (3 M), and in the Perris Collection in Montpellier (1 M, 3 F).

Identity: Aritranis mediterraneus (Tschek,1872). New synonym.

Ichneumon odoriferator Dufour et Perris

Ichneumon odoriferator Dufour et Perris, 1840: 44. Lectotype hereby

designated: M, "1790. M", "Dufour collectio". Paris Museum, found in the

Giraud collection, and transferred to the general collection under Hoplocrypus

graefel,

Identity: Aritranis odoriferator (Dufour et Perris,1840), syn.

Hoplocryptus graefei Thomson, 1896. New synonym.

Ichneumon odynericidus Dufour et Perris

Ichneumon odynericidus Dufour et Perris, 1840: 45. Lectotype hereby

designated: F, "5691'"'", "Museum Paris France Dufour". Paris Museum, found

in the Dufour Collection, and transferred to the general collection under

Kaltenbachia bimaculata.

Identity: Xylophrurus augustus (Dalman,1823) (Giraud 1866: 480, Thomson

1888: 1237, Schmiedeknecht 1890: 118).

Anomalon mandibulator Dufour et Perris

Anomalon mandibulator Dufour et Perris, 1840: 46. Types lost.

Identity: Isadelphus armatus (Gravenhorst,1829). New synonym. Inter-

preted in accordance with Giraud (1866: 484).

Pimpla ephippiatoria Dufour et Perris

Pimpla ephippiatoria Dufour et Perris, 1840: 47. Lectotype hereby

designated: F, "3266. F", "Museum Paris France Dufour". Paris Museum,

found in the Dufour Collection, and transferred to the general collection under

Perithous divinator.

Identity: Perithous divinator (Rossi,1790) (Giraud 1866: 486).

Horstmann: Ichneumonidae 105

Pimpla marginellatoria Dufour et Perris

Pimpla marginellatoria Dufour et Perris, 1840: 48. Types lost.

Identity: Perithous septemcinctorius (Thunberg 1822). New synonym.

The male mentioned by Giraud (1866: 487) is still preserved in the Perris col-

lection. It is not a syntype of Pimpla marginellatoria, and it belongs to the

species Pseudopimpla pygidiator Seyrig, 1927.

BOYER DE FONSCOLOMBE

The following notes are a supplement to my revision of the species of

Ichneumonidae described by Boyer de Fonscolombe (Horstmann 1980).

Ichneumon insidiator Boyer de Fonscolombe

Ichneumon insidiator Boyer de Fonscolombe, 1847 in 1847-1854: 66.

Holotype: M, "28. I. insidiator nob." (blue label) (the head, and one hind

wing lacking).

Identity: Spilothyrateles fabricii (Schrank,1802). I had identified

Ichneumon insidiator as Amblyteles truncicola Thomson, 1888 (Horstmann

1980: 131), but had overlooked, that the latter species is considered to be a

synonym of Spilothyrateles fabricii .

Phygadeuon incertus Boyer de Fonscolombe

Phygadeuon incertus Boyer de Fonscolombe, 1851 in 1847-1854: 112. Holo-

type: M, "8. Ph. incertus nob." (Blue label), "Museum Paris 1867 Coll.

O. Sichel". Paris Museum, found in the de Gaulle Collection, and transferred

to the box with types of species described by Boyer de Fonscolombe in the

general collection.

Identity: Phygadeuon incertus Boyer de Fonscolombe, 1851.

LUCAS

H. Lucas (1849) described 14 new species of Ichneumonidae, the types of

which he had collected in Algeria. The material from Algeria is preserved in

the Paris Museum as a separate Collection. Especially the Hymenoptera Para-

sitica of that collection are in a bad order, and partly destroyed by Dermesti-

dae. Therefore, I transferred all types of Ichneumonidae to a special box in

the general collection. The typical labels of the Lucas material are green

round labels with numbers, rarely with locality names on the underside, and

rectangular name labels. The meaning of those numbers is not known to me.

Pimpla nigrolineata Lucas

Pimpla? nigrolineata Lucas, 1849: 323. Lectotype hereby designated:

F, "689.", "Pimpla?nigrolineata, Luc. F".

Identity: Odinophora nigrolineata (Lucas 1849), syn. Pimpla perezi

Kriechbaumer, 1898. New synonym.

106 Contrib. Aner. ents Inst. vol. 20, 1983

Glypta erythrogaster Lucas

Glypta erythrogaster Lucas, 1849: 324. Lectotype hereby designated:

M, "689.", "'Glypta erythrogaster, Luc. M". A female syntype is present, but

partly destroyed.

Identity: Glypta cylindrator (Fabricius,1787) (syn. Glypta bicornis Boie

1850). New synonym. |

Lissonota flavipes Lucas

Lissonata flavipes Lucas, 1849: 325. Lectotype (? holotype) hereby

designated: M, "Oran", "Lissonota flavipes, Luc. M".

Identity: Lissonota flavipes Lucas, 1849. I was not able to identify

the type with a described species.

Bassus flavipes Lucas

Bassus flavipes Lucas, 1849: 325. Lectotype (? holotype) hereby

designated: M, "949" (?), "Bassus flavipes, Luc.".

Identity: Sussaba flavipes flavipes (Lucas, 1849), syn. Sussaba

coriacea neopulchella Diller, 1980 (syn. Bassus pulchellus pulchellus auct.).

New synonym.

Anomalon affine Lucas

Anomalon affine Lucas, 1849: 327. Lectotype labelled by Schnee and

hereby designated: M, "689", "Anomalon affine , Luc. M".

Identity: Barylypa delictor (Thunberg,1822) (Schnee det.). New

synonym.

Collyria erythrogaster Lucas

Collyria erythrogaster Lucas, 1849: 328. Lectotype (? holotype) here-

by designated: M, "1291", "Collyria erythrogaster, Luc. M".

Identity: Phaenolobus fulvicornis (Gravenhorst,1829) (Hinz det.). New

synonym.

Collyria armata Lucas

Collyria armata Lucas, 1849: 328. Lectotype hereby designated: M,

"689", "Collyria armata, Luc. ?M".

Identity: Pristomerus armatus (Lucas 1849), syn. Pristomerus orbitalis

Holmgren, 1860. New synonym. Pristomerus gratiosus Tosquinet, 1896, also

described from Algeria, is considered to be an African subspecies of

Pristomerus orbitalis, differing in color from the European subspecies

(Aubert 1968a: 142). The lectotype of Collyriaarmata , however, does not dif-

fer in color from European material of the species.

Cryptus Levaillantii Lucas

Cryptus Levaillantii Lucas, 1849: 329. MHolotypes: F, "Oran",

Cryptus Levaillantii, Luc.".

Identity: Synechocryptus levaillantii (Lucas,1849) (van Rossem 1969b:

371).

107

Horstmann: Ichneumonidae

Cryptus bicolor Lucas

Cryptus bicolor Lucas, 1849: 330. Lectotype designated by van Rossem

(1969b:: 358): 7, "290" “crupcusybrcolom Lac.’, |

Identity: Trychosis legator (Thunberg,1822) (van Rossem 1969b: 358).

Cryptus armatus Lucas

Cryptus armatus Lucas, 1849: 330. Lectotype hereby designated: F,

"6890" Croptustarmatus,: Luciah!’.

Identity: Meringopus armatus (Lucas 1849). The species is similar to

Meringopus turanus (Habermehl 1918) (see van Rossem 1969a: 179), but differs

from the latter in the following characters: Body size 13-l4 mm. Antennae

with 41-42 segments, the 18th segment about 1,3 times as long as broad.

Speculum punctate. Ovipositor sheaths a little shorter than the hind tibiae

(Female of Meringopusturanus: Body size 15-16 mm. Antennae with 55 seg-

ments, the segments in the apical third more slender. Speculum polished, at

least partly. Ovipositor sheaths a little longer than the hind tibiae).

Cryptus annulicornis Lucas

Cryptus annulicornis Lucas, 1849: 33l. Lectotype designated by

van Rossem (1969b: 370): F, "490.", "Cryptus annulicornis, Luc." (both tips

of antennae, both front legs, one middle and one hind leg, and the greater

part of the gaster lacking).

Identity: Nomen oblitum (van Rossem 1969b: 370).

Ichneumon rubrocinctus Lucas

Ichneumon rubrocinctus Lucas, 1849: 333. Holotype: F (!), "299.",

"Tchneumon rubrocinctus, Luc. M ()).

Identity: Triptognathus rubrocinctus (Lucas,1849), syn. Amblyteles

impolitus Berthoumieu, 1894. New synonym.

Ichneumon flavomaculatus Lucas

Ichneumon flavomaculatus Lucas, 1849: 333. Holotype: M, "689.",

"Ichneumon flavomaculatus, Luc. M".

| Identity: Ichneumon (s. 1.) flavomaculatus Lucas, 1849. I was not able

to identify the type with a described species.

Mesoleptus flavovariegatus Lucas

Mesoleptus flavovariegatus Lucas, 1849: 334. Lectotype (? holotype)

hereby designated: M, "689", "Mesoleptus flavovariegatus, Luc." (both

antennae, both middle legs, and the greater part of the gaster lacking, the rest

partly destroyed by Dermestidae).

Identity: Lissonota lineata (Gravenhorst 1829). New synonym.

GIRAUD

Giraud (1857-1872) described 19 new European species of Ichneumonidae.

His collection is preserved in the Paris Museum, but some material had been

transferred to the general collection of Ichneumonidae of that museum. The

108 Contribs. Amer. Ent..Inst... vol, 20, 1983

types of some species should be in the Museum of Natural History in Vienna,

but could not be found there (Fischer, in lit.). Some of those types, however,

now are in the Paris Museum too. Perhaps Giraud took them with him, as he

moved from Vienna to Paris. In many cases the material collected by Giraud

is imperfectly labelled. He often used labels with only collecting dates or

abbreviations of the locality or the host, and often labelled only the first speci-

men of a series.

Metopius nasutus Giraud

Metopius nasutus Giraud, 1857: 169. Lectotype hereby designated: F,

"Ullr." (? = Ullrich, name of a collector), "F", "Museum Paris 1877 Coll.

Giraud". Paris Museum, general collection under Peltopius vespoides.

Identity: Metopius vespoides (Scopoli 1763) (Rogenhofer and Dalla Torre,

1882: 600).

Mesostenus nubeculator Giraud

Mesostenus nubeculator Giraud, 1857: 170. Lectotype hereby designated:

F, "Ill fig. G" (referring to a figure in the original description), "Museum Paris

1877 Coll. Giraud". Paris Museum, general collection under Listrognathus

pubescens. ?

Identity: Listrognathus pubescens (Boyer de Fonscolombe,1850). New

synonym.

Anomalon fasciatum Giraud

Anomailon fasciatum Giraud, 1857: 170. Lectotype hereby designated: :

F, "Sicil.", "Museum Paris 1877 Coll. Giraud". Paris Museum, general collec-

tion under Gravenhorstia picta.

Identity: Gravenhorstiapicta Boie, 1836 (Bridgman and Fitch, 1884: 228).

Pezomachus tricolor Giraud

Pezomachus tricolor Giraud, 1857: 171. Lectotype hereby designated:

F, "Prat. avr.". Paris Museum, Giraud Collection, box 31.

Identity: Gelis cyanurus (Forster, 1850). New synonym.

Ichneumon amabilis Giraud

Ichneumon amabilis Giraud, 1863a: 20. Primary homonym of Ichneumon

(Exephanes ?) amabilis Kriechbaumer, 1895. Lectotype hereby designated:

F, “Susa", "Museum Paris 1877 Coll. Giraud". Paris Museum, general collec-

tion under Ichneumon lautatorius .

Identity: Ichneumon lautatorius Desvignes, 1856. New synonym.

Hemiteles Liparae Giraud

Hemiteles Liparae Giraud, 1863b: 1267. Holotype: M, "ex Lipara

tomentosa 29 mars.", ''Hemiteles Liparae Giraud". Paris Museum, Giraud

Collection, box 31.

Identity: Gelis liparae (Giraud,1863).

109

Horstmann: Ichneumonidae

Pimpla cheloniae Giraud

Pimpla cheloniae Giraud, 1869: 149. Lectotype hereby designated: F,

"cervini 20 mai", "de Chelonia cervini (Valais, Fallou)" (the latter label

refers to nine syntypes). Paris Museum, general collection under Pimpla.

Identity: Pimpla cheloniae Giraud, 1869.

Ischnus tineidarum Giraud

Ischnus tineidarum Giraud, 1872a: 393. Lectotype hereby designated:

F, "Fallou", 'Talaeporia Pseudobombycella St. Germain 29 Avril". Paris

Museum, Giraud Collection, box 26.

Identity: Rhexidermus tineidarum (Giraud, 1872).

Ischnus nigrinus Giraud

Ischnus nigrinus Giraud, 1872a: 395. Lectotype hereby designated:

F, 'de Coleoph. Giraudi (Oisans)", "Ischnus nigrinus m.". Paris Museum,

Giraud Collection, box 26.

Identity: Rhexidermus nigrinus (Giraud, 1872).

Plectiscus tenthredinarum Giraud

Plectiscus tenthredinarum Giraud, 1872a: 396. Lectotype hereby desig-

nated: F, "Vine 13 mai", "Plectiscus tenthredinarum m.". Paris Museum,

Giraud Collection, box 39.

Identity: Adelognathus tenthredinarum (Giraud 1872), syn. Adelognathus

nigricornis Thomson, 1888 (following the interpretation of Perkins, 1943b, the

types of Adelognathus nigricornis being lost) (Hinz vid.). New synonym.

Roman (1918: 19) erroneously identified Adelognathus tenthredinarum as

Adelognathus pusullus Holmgren, 1857.

Perilissus luteocephalus Giraud

Perilissus luteocephalus Giraud, 1872a: 397. Lectotype hereby desig-

nated: F, "in copula" (labels of the corresponding male: "Grenoble 23 avril

in copula", "de Selandria micans K1.","Perilissus luteocephalus Giraud").

Paris Museum, Giraud Collection, box 41.

Identity: Perilissus tripunctator (Thunberg,1822) (syn. Perilissus

longicornis Brischke, 1871). New synonym.

Tryphon lateralis Giraud

Tryphon lateralis Giraud, 1872a: 401. Primary homonym of Tryphon

lateralis Gravenhorst, 1829. Lectotype hereby designated: F, "Vine 19 mai".

Paris Museum, Giraud Collection, box 43.

Identity: Monoblastus caudatus Hartig, 1837. New synonym.

Trematopygus selandrivorus Giraud

frematopygus selandrivorus Giraud, 1872a; 402. Lectotype labelled by

Aubert and designated by Hinz (1975: 45): M, "I7 avr.", "g. c." (2),

"Trematopygus selandrivorus m.". Paris Museum, Giraud Collection, box 43.

110 Contrib. Amer. Ent. Inst., vol. 20, 1983

Identity: Glyptorhaestus selandrivorus (Giraud, 1872) (Hinz 1975: 45).

Trematopygus aprilinus Giraud

Trematopygus aprilinus Giraud, 1872a: 403. Lectotype labelled by

Aubert and hereby designated: F, "De Selandr. melanoceph. 18 avr.",

"Trematopygus aprilinus m.". Paris Museum, Giraud Collection, box 43.

Identity: Trematopygodes aprilinus (Giraud , 1872) (Aubert 1968b: 69).

Polyblastus annulicornis Giraud

Polyblastus annulicornis Giraud, 1872a: 406. Holotype: F, "Prater

13 sept.", "Polyblastus annulicornis Giraud F.". Paris Museum, Giraud Col-

lection, box 43.

Identity: Ctenochira grossa (Brischke 1871) (Schmiedeknecht 1912 in 19ll-

1927: 2466). As the papers of Brischke and Giraud are dated both from 1871,

it was necessary to look for further information about the dates of publication.

The volume XI of the "Schriften der koniglichen physikalisch-okonomischen

Gesellschaft Konigsberg", which contains Brischke's paper, arrived at Vienna

during the week before January 4, 1872 (Sitzungsberichte der mathematisch-

naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften,

Wien, 65, Abt. I, 1872: 4). The third and fourth parts of the volume (V) I of

the "Annales de la Societe entomogique de France", which contain Giraud's

paper, came out on February 14, 1872, and on April 10, 1872, respectively, and

arrived at Bruxelles in March and in May of that year (Annales de la Societe

entomologique de Belgique, Comptes Rendus des Seances de la Societe entomol-

ogique de Belgique 15, 1872: LI and LXIX).

Echthrus populneus Giraud

Echthrus populneus Giraud, 1872a: 407. Holotype: F, "de Saperda

populnea 23 avr.", "Echthrus populneus Giraud". Paris Museum, Giraud Collec-

tion, DOx Si.

Identity: Schreineria annulata (Brischke, 1864). New synonym.

Diplomorphus thoracicus Giraud

Diplomorphus thoracicus Giraud, 1872a: 409. Lectotype hereby deésig-

nated: F, "Vallouise h.-alp.", "Diplomorphus thoracicus Giraud". Paris

Museum, Giraud Collection, box 43.

Identity: Boethus thoracicus (Giraud,1872) (Schmiedeknecht 1912 in 191l-

1927: 2396).

Aglyphus nigripennis Giraud

Aglyphus nigripennis Giraud, 1872a: 4ll. Lectotype hereby designated:

F, "Bolo." (= Bologne), "Aglyphus bicolor Giraud." CQ), “Diadochus nN, gen.

bicolor m. F." (!). Paris Museum, Giraud Collection, box 43.

Identity: Scolobates nigripennis Sichel, 1860 (de Gaulle,1908: 60).

Pimpla gallicola Giraud

Pimpla gallicola Giraud, 1872b: 506. Lectotype hereby designated:

F, "Andr. gland. avr." ( = Andricus glandulae Hartig , 1840). Paris Museum,

Giraud Collection, box 48.

Horstmann: Ichneumonidae +71

Identity: Scambus planatus (Hartig,1838). New synonym. Perkins (1943a:

267) and subsequent authors incorrectly synonymized Pimpla gallicola Giraud

with Scambus vesicarius (Ratzeburg 1844). There is a female of the latter

species in the Giraud collection, which had been bred from "wematus

gallarum" ( = Pontaniaviminalis Linnaeus, 1858), but it cannot be eee

as a syntype of the Giraud species, for only. oes are mentioned in the origin-

al description.

At the end of his life, Giraud gave a list of parasites and the correspond-

ing hosts to A. Laboulbene for publication, for he himself was no longer able

to make it ready for print. Laboubene (1877) added some footnotes with re-

marks on the biology of the host species, and published that list after Giraud's

death. The list contains 33 names of ichneumonid species, which are attributed

to Giraud as the author, but were not described by him. Three of those cases

must be discussed below, for in the list some additional notes are given, which

might be considered as indications in the sense of article 16 of the Rules of

Nomenclature.

Amblyteles montivagus Giraud

Amblyteles montivagus Giraud in Laboulbene, 1877: 399. Following the

text in the list ("Amblyteles montivagus Giraud, A. fossorius V. I. Grav."),

Amblyteles montivagus Giraud is considered to be a new name for Amblyteles

fossorius Var. 1 sensu Gravenhorst (1829: 165) (Article 16, a, i of the Rules).

As the notes in the list are from Giraud, the name is attributed to him. The

name is a primary homonym of Amblyteles montivagus Berthoumieu, 1896.

Lectotype hereby designated: F, "f.". Museum of Natural History, Wroclaw,

Gravenhorst collection under Ichneumon fossorius var. |.

Identity: Ctenichneumon messorius (Gravenhorst, 1829) (Hinz vid.). New

synonym.

Pezomachus glallarium Giraud

Pezomachus gallarum Giraud in Laboulbene, 1877: 402. As it is not pos-

sible to interpret unequivocally the text in the list ("Pezomachus gallarum

Giraud (M Hemit. bicolor, var. 4)"), the name is considered to be a nomen

nudum.

Pimpla gallarum Giraud

Pimpla gallarum Giraud in Laboulbene, 1877: 409. It is clear from the

text in the list ("Pimplagallarum Giraud, stercorator var. Grav.") and from

the published host species, that the name is a lapsus for Pimpla gallicola

Giraud, 1872.

LIST OF NOMINA NUDA

Ichneumon plusiae Giraud in Laboulbene, 1877: 398

Ichneumon valaisianus Giraud in Laboulbene, 1877: 399

Hoplismenus nigripes Giraud in Laboulbene, 1877: 399

Trogus Bellieri Giraud in Laboulbene, 1877: 399

Phaeogenes ruficornis Giraud in ED ubaaibene, 1877: 400

Cryptus leucostilus Giraud in Laboulbene, 1877: 400

Cryptus zygaenae Giraud in Laboulbene, 1877: 40]

112 Contrib. Amer. Ent. Inst., vol. 20, 1983

Hemiteles aranearum Giraud in Laboulbene, 1877: 401

Hemiteles brevicornis Giraud in Laboulbene, 1877: 401

Hemiteles scutellaris Giraud in Laboulbene, 1877: 402

Pezomachus coleophorae Giraud in Laboulbene, 1877: 402

Pezomachus diastrophi Giraud in Laboulbéne, 1877: 402

Pezomachus gallarum Giraud in Laboulbene, 1877: 402

Pezomachus palustris Giraud in Laboulbene, 1877: 402

Pezomachus potentillae Giraud in Laboulbene, 1877: 402

Porizon perlae Giraud in Laboulbene, 1877: 403 |

Campoplex coleophorae Giraud in Laboulbene, 1877: 404

Campoplex curtus Giraud in Laboulbéne, 1877: 404

Campoplex helicinellae Giraud in Laboulbéne, 1877: 404

Campoplex setinae Giraud in Laboulbéne, 1877: 404

Campoplex transfuga Giraud in Laboulbéne, 1877: 405

Anomalon proteae Giraud in Laboulbene, 1877: 405

Anomalon zygaenae Giraud in Laboulbéne, 1877: 405

Mesochorus falcifer Giraud in Laboulbene, 1877: 406

Mesochorus fusculus Giraud in Laboulbéne, 1877: 406

Mesochorus sulcatus Giraud in Laboulbéne, 1877: 406

Mesochorus tachinarum Giraud in Laboulbéne, 1877: 407

Bassus pipizae Giraud in Laboulbene, 1877: 408

Lissonota pugnator Giraud in Laboulbene, 1877: 408

Pimpla orgyiae Giraud in Laboulbéne, 1877: 409

Pimpla palliata Giraud in Laboulbéne, 1877: 409

LABOULBENE

A. Laboulbene (1858) described one new species of French Ichneumonidae,

which was bred from a spider. The types were found in the Fairmaire collec-

tion in the Paris Museum.

Pimpla Fairmairii Laboulbene

Pimpla Fairmairii Laboulbene, 1858: 814. Lectotype hereby designated:

F, "Pimpla Fairmairii A, S. E. Fr." (= Ann. Soc. Ent. France) (partly

destroyed by Dermestidae, one front and one hind wing lacking). Paris Museum,

found in the Fairmaire Collection, and transferred to the general collection

under Zaglyptus multicolor,

Identity: Zaglyptus multicolor (Gravenhorst,1829) (Morley 1908: 124).

SICHEL

J. Sichel (1860) described one new species of Ichneumonidae from Sicily.

The type is preserved in his collection in the Paris Museum.

Scolobates nigripennis Sichel

Scolobates nigripennis Sichel, 1860: 758. Holotype: F (!), "Sic 59"

(= Sicile), "Scolobatesnigripenis ... n.s." (!, partly illegible). Paris Museum,

Sichel Collection.

Identity: Scolobates nigripennis Sichel, 1860 (de Gaulle 1908: 60).

Horstmann: Ichneumonidae 113

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Mexico: 1. Subfamily Metopiinae. Bull. U. S. Natl. Mus. 216: 1-318.

A NEW EASTERN PALEARCTIC GENUS OF THE

SUBFAMILY PHRUDINAE

(HYMENOPTERA : ICHNEUMONIDAE)

D. R. Kasparyan

Zoological Institute, Academy of Sciences

Leningrad, V-l64, USSR

A new genus, Lygurus and a new species, L. townesi, is described from

Khabarovsk region. It is closely related to the Ethiopian Erythrodolius

Seyrig (Townes 1971: 32), but differs in the structure of the frons, clypeus,

and more elongate maxilla and labium.

LYGURUS, new genus

Type-species: Lygurus townesi, n. sp.

Front wing 4.3 mm. Frons without a median longitudinal carina, with

small cavities upper of each antennal socket (Fig. 3). Cheek without a

vertical groove. Clypeus broad, centrally with weak transverse ridge; its api-

cal margin blunt, more sharp laterally. Lower tooth of mandible scarcely

shorter than upper tooth. Maxilla and labium elongate, projecting below apex

of clypeus by about one third the height of head; the maxillary palpus as

long as height of head. Mesopleuron polished, with coarse puncture along its

front margin, under subtegular carina, and in its lower half. Propodeum

areolated (Fig. 4). Apex of front tibia with a scarcely visible tooth on outer

side. Tarsal claws simple, with 1-2 teeth basally. Wing venation as figured

(Fig. 1). First tergite with strong glymma. Tergite 2 polished, basolateral

with short and sharp groove. Hairs on tergite 2 very short and slender, but

rather dense. Epipleuron of tergite 2 separated by crease, bent under and

overlieing. Epipleuron of tergite 3 wide bent but not separated by crease

(with weak crease in basal 0.2). Ovipositor sheath 1.2 as long as body. Ovi-

positor slender, cylindric, in apical third strongly upcurved and somewhat de-

pressed.

The generic name is from the Greek lygos (flexible switch) plus ura

(tail), referring to long flexible ovipositor.

One Palearctic species is known, collected in taiga near Khabarovsk. The

The species is described below.

Lygurus townesi, n. sp. (Figs. 1-4 )

Female (holotype): Length 5.7 mm. Ovipositor 7 mm. Flagellum 22-

segmented. Scape subcylindric, elongated, with apical truncation approximate-

ly transverse. Head strongly narrowed behind eyes (Fig. 2). Cheek 0.8 as

long as basal wide of mandible. Clypeus on apical margin with long hairs.

Occipital carina joining hypostomal carina at a considerable distance above

base of mandible. Pronotum with weak epomia. Notauli very short. Scutel-

lum with distinct lateral carinae from its base to apex. Propodeum short,

completely areolated (Fig. 4). Tibiae beside usual short hairs on outer side

with numerous more strong bristles. First tergite strongly elongated, 3 times

as long as apical wide (Fig. 4); its dorsolateral carinae present (some weaker

behind the spiracles), dorsal longitudinal carinae almost obliterated. Glymmae

large and deep, separated by a translucent partition. Abdomen behind the

Kasparyan: Phrudinae

Figs. 1-4, Lygurus townesi, gen. et sp. n.: lL,

insect. 2, Head, dorsal view.

first two tergites.

3, Head, frontal view.

Side view of whole

4, Propodeum and

Vis Gontrib. Amer Ent.‘Inst., vol. 20, 1983

middle of tergite 3 strongly compressed. Other structure as figured, and as

described under generic heading.

Face, frons, clypeus basally, pronotum laterally, mesonotum, scutellum,

metapleuron with moderately small and dense punctures. Vertex, temple, and

first lateral areas of propodeum very finely punctate. Mesopleuron polished,

its front and upper margin and its lower half with rather large punctures that

are separated by about 1-2 their diameter.

Coloration. Black. Antennae dark brown. Scape below, upper margin

of pedicel, frontal and vertical orbits, cheek, lower part of temple adjoining

the mandible yellowish brown; facial orbits yellow. Face, temples, base of

clypeus, anterior and posterior margins of lateral parts of pronotum, subtegu-

lar ridge, epimeron greyish-brown. Clypeus apically, mandible (except apices),

maxilla and labium, fore and middle legs reddish-yellow; tegulae reddish-brown.

Hind legs brownish with reddish-yellow trochantelli, base of femora and inner

side of tibia; tibia with basal black spot. Tergite 2 with reddish hind margin.

Pterostigma blackish, basally with small white spot.

Male: Unknown.

Holotype: Female, Ridge Khekhcir near Khabarovsk, taiga, 22.VII.1981.

(Kasparyan). In Zoological Institute, Leningrad.

RELATIONSHIPS AND TAXONOMIC PLACEMENT OF THE

GENUS sBLacus (HYMENOPTERA: BRACONIDAE)

M. Capek

Forest Research Institute, Zvolen,

Research Station Banska Stiavnica, Czechoslovakia

The genus Blacus Nees von Esenbeck, 1818, is one of the large Braconid

genera, not only by the number of known species (Haeselbarth 1973 lists 43

species from Europe and Central Asia in his revision) but also by the abundance

of some of them. They are often observed swarming in large numbers like

the Chironomidae. Information on their host relations, biology, and develop-

mental stages is contradictory or scanty. The classification of the genus is

chiefly based on external morphology of the adults. Consequently there are

often disagreeing opinions on the relationships of the genus and its position

in the Braconid hierarchy.

3 HOST RELATIONS OF BLACUS SPECIES

There are several references on the hosts of Blacus species, but most

of them are unreliable. The first report of Bouché (1834) refers to Lacanobia

oleracea as the host of Blacus humilis. This is almost certainly erroneous,

and probably pertains to an Apanteles sp. The fore wing venation of

Apanteles resembles to a certain extent that of Blacus. All other records 7

of lepidopteran hosts are also unlikely, like Mamestra brassicae for B. humilis

(Rondani, 1876), Tortrix sp. and Bucculatrix nigricomella for B. maculipes

(de Gaulle, 1907), Simyraalbovenosa for B. sp. (Cole, 1931), Tortrix

viridiana and Rhyacionia buoliana for B. sp. (Kolubajiv, 1962a), Coleophora

conspicuella for B. sp. (Kolubajiv, 1962b), and Thaumetopoea pityocampa for

B. instabilis (Anonymous, 1963). There is little doubt that three records

of Hymenoptera and one of Aphidoidea as the hosts of Blacus species are also

incorrect: Nematus sp. for B. errans (Rudow, 1918), Megastigmus brevicaudis

for B. exilis (Ahlberg, 1927), Spilomena troglodytes for B. longipennis ;

(Schutze et Roman, 1931), and Phorodon cannabis for B. ruficornis (Goidanich,

1928).

Several dipteran hosts also appear in the literature for Blacus species.

Goureau (1851) reported Agromyza nana as the host of B. ruficornis and from

‘the same host he described B. florus Rondani (1876) described B. brachialis

from Chlorops lineatus. Further dipterous host records are Fungivora

signata as the host of B.maculipes (Fahringer, 1922), Philophylla heraclei

for B. exilis (Lundblad et Lindblom, 1925), and Rhabdophaga saliciperda

for B. errans (Telenga, 1935). These host species develop in plant tissues

(galls, mines, fungi, etc.) where other insects often live. This increases the

probability of associating Blacus species with incorrect hosts, especially in mass

rearings. The most recent host records are of Oscinella frit for B. inter-

stitialis (Fischer, 1963), Contarinia medicaginis for B. instabilis (Anony-

mous, 1963), Acanthiophilus helianthi for B. ? impudians (Fulmek, 1968),

and Contarinia medicaginis for B. exilis (Haeselbarth, 1973).

The coleopterous hosts of Blacus species are mentioned in the literature

most often and seem to be the most reliable, although there are obviously some

mistakes. Boudier (1834), described B. barynoti from Barynotus elevatus

119

120 Contrib. Amer. Ent. Inst., vol. 20, 1983

Ratzeburg (1848) mentioned Stereonychus fraxini as the host of B. hastatus.

Ruthe (1861) recorded Scolytus mali, S. rugulosus, Stenostola nigripes,

Tetrops praeusta, Pogonochaerus hispidus, and Dasytes plumbeus, as possible

hosts of B. errans. Goureau (1862) described B. fuscipes from Scolytus

rugulosus. Further references are found in the host-parasite catalogues or

parasite lists of the following authors: de Gaulle (1907) mentioned Barynotus

obscurus and Otiorrhynchus ligneus as hosts of B. barynoti, de Gaulle (1908)

Exocentrus adspersus for B. errans and Barynotus obscurus for B. pallipes,

Wahl (1914) Anisandrus dispar and Xyleborinus saxeseni for B. fuscipes ,

Feytaud (1914) Otiorrhynchus spp. for B. pallipes, Rudow (1918) Barynotus

sp. and Miaruscampanulae for B. armatulus, and B. humilis, Otiorrhynchus

sp., and Barynotus sp. for B. barynoti , Gymnetron sp. and Orchestes sp. for

B. conformis, Liophloeus schmidti for B. diversicornis , Ceutorrhynchus sp.

and Apoderuscoryli for B. maculipes, Curculio glandium for B. tripudians,

and Cionusthapsi for B. humilis Ns., Schutze et Roman (1931) Dasytes

coeruleus for Blacus sp., Telenga (1935) Cryptocephalus lycoperdi for B.

humilis, Morley (1937) cis sp. for B. hastatus, Thompson (1953) Otiorrhynchus

rhynchus laevigatus for B. barynoti, and Pityogeneschalcographus for B.

fuscipes, Wayman (1957) Cryptophagus subfumatus for B. instabilis,

Jourdheuil (1960) Meligethes aeneus for B. hastatus, Kolubajiv (1962a)

Magdalis violacea, Pissodes harcyniae, Dendroctonusmicans , and Ernobius

pini for Blacus sp., Lipkow (1965) Tachyporus obtusus for B. ruficornis ,

and Alford (1975) Antherophagus sp. from bumblebee nests for B. paganus .

The more reliable records are of Fischer (1962) of B. koenigi from

Scolytus koenigi. Valuable data on host associations are in Haeselbarth's

revision of the genus: Ceutorrhynchus sp. and Meligethes aeneus for B.

nigricornis, Magdalisarmigere and Pityokteines vorontzowi for B. exilis ,

Dryocoetes autographus, Abdera flexuosa, and Triplax russica for B.

hastatus, Blastophagus piniperda and Sitodrepa panicea for B. humilis , and

Cryptophagus sp. and Carpophilus sp. for B. instabilis. A total of 46

species of Coleoptera (Curculionidae 17, Ipidae 10, Cerambycidae 5, Nitidulidae

and Cryptophagidae 3, Cantharidae and Anobiidae 2, and Erotylidae, Cisidae,

Melandryidae, and Staphylinidae 1 species) are mentioned in the liaterature as

the hosts of various Blacus species.

Among the Braconids that I have received from many entomologists dur-

ing the past 25 years for naming, there were only two specimens of Blacus

reared from the known hosts. The first was B. errans Ns. reared by Profes-

sor Capecki of Cracow, Poland from Dasytes sp., while the second was

B. longipennis Grav. reared by Dr. Kinelski of Warsaw, Poland from Anaspis

sp.

The species of Blacus seem to have specialized as the parasites of the

Coleoptera and less probably of the Diptera. The host records from other

insect orders are evidently erroneous, and are misidentifications, either of the

hosts or the parasites.

BIOLOGY OF BLACUS SPECIES

There is even less reliable information available on the biology of Blacus

species. Bouche (1834) stated that B. humilis Ns. was a gregarious

Capek: Blacus 23

endoparasite in the larvae of Lacanobia oleracea and pupated in a loose heap

of greyish cocoons in the host pupal cell in the soil. This report seems to

be rather doubtful and most probably it relates actually to an Apanteles

species. Several authors further recorded Blacus species as internal parasites

of the host larvae. Only Lipkow (1965) B. ruficornis as an internal parasite

of the host adult and Rondani (1876) as an internal parasite of the host pupa.

Although Stich (1929) stated that Blacus species are parasites of the adult

Coleoptera, the published evidences indicate that they are larval parasites.

This is confirmed by my observations that the remains of the coleopterous

larvae were attached to the cocoons from which B. errans and B. longipen-

nis emerged.

The dancing or swarming habit of several Blacus species has been long

known. More recently Haeselbarth (1973) refers to the swarming of B.

maculipes, B. nigricornis, and B. longipennis . Most authors observed only

males in the swarms, but Smits van Burget (1921) observed both sexes of

B. ruficornis. The swarming represents the nuptial flight (konig, 1964; Tobias,

1965; and Gruhl, 1967). Blacus spp. are often captured by the light traps.

The nocturnal activity is typical for them (Tobias, 1965; Haeselbarth, 1973;

personal observation).

The number of generations of B. hastatus was studied by Jourdheuil

(1960) and of B. ruficornis by Konig (1967). Both species are bivoltine or

polyvoltine. Smits van Burgst (1921) and Hancock (1925) observed that only

the females of B. ruficornis overwinters.

It therefore appears that the species of Blacus are solitary endoparasites

of the host larvae. Most of them are nocturnal and swarming congregation

of the males is typical.

TAXONOMY OF LAST INSTAR LARVAE OF BLACUS SPECIES

, Up to present time only the larva of B. longipennis (as Blacus sp. in

Capek, 1970) was described and figures (Fig. 1). Now the last instar larval

cast skin of B. errans has been studied. (Fig. 2). On the base of these two

species the larvae of Blacus spp. can be characterized as follows: mandibule

with more or less distinct base, the blades with two rows of 4-5 teeth; labial

sclerite almost rectangular, basically round; labial palpi with one large sensil-

lum and 1-2 small sensilla; stipital sclerite and cardo distinct; hypostoma rudi-

mentary, hypostomal spur indicated as well as epistoma; antenna disc-shaped,

with visible sensilla; spiracle with cup-shaped atrium, which is about three

times as wide as the closing apparatus; skin with two types of spines—-most

spines are of about one sixth of mandible length, a few longer spines are about

half as long as mandible.

The generalized form of most head sclerites in the larvae of Blacus

species resemble the Braconid genera like Zubazus, Triaspis, Aliolus and

Allodorus.

RELATIONSHIPS OF BLACUS TO OTHER BRACONID GENERA

Blacus was originally placed between Microgaster and Rogas (= Macro-

centrus) by Nees von Esenbeck (1818) Other authors placed it variously, as:

20,

Contrib. Amer. Ent. Inst., vo

Fae

> Z

Sun

acG

CO tm

den)

n ©

s 8

bas

Cpe

woe

Psat oe

a =

17 p)

(40)

ro)

Figs. 1-2.

1, Blacus longipennis Grave

Capek: Blacus t23

Haliday (1835) between Perilitus and Helcon, Wesmael (1835) between

Hybrizon and Leiophron, and Ratzeburg (1852) between Pleiomerus and

Eubadizon. Foerster (1862), Marshall (1889), Dalla Torre (1898), Schmiedeknecht

(1930), Watanabe (1937), and Docavo Alberti (1960) placed Blacoidae between

Brachistoidae and Liophronoidae, Ashmead (1900) between Helconinae and

Sigalphinae, Szépligeti (1904) and Viereck (1918) between Liophroninae and

Ichneutinae, Granger (1949) between Meteorinae and Calyptinae, and Muesebeck

and Walkley (1951) between Helconinae and Agathidinae. Boutek and Stary

(1957), Capek (1969), and Achterberg (1976) placed it in Helconinae. Tobias

(1965), Shenefelt (1969), and Mason (1974) placed it in Euphorinae, and finally

Marsh (1979) in Blacinae.

Most recent authors place Blacus in the tribe Centistini of the subfamily

Euphorinae (Pygostolus, Centistes, Syrrhizus, Allurus). The tribe is rather

homogenous by its host relations (Coleoptera), biology, (parasites of the adults)

as well as by its larval taxonomy (Capek 1970). Their females are character-

ized by having a down curved strong ovipositor well adapted for oviposition into

hardly sclerotized coleopterous adults. In all these characters Blacus differs

from Centistini and thus its placement in the Euphorinae is unjustified. On

the contrary, Blacus exhibits most of the characters of the tribe Brachistini

(sensu Mason 1974) of the subfamily Helconinae.

REFERENCES

For references not included here, see Shenefelt, 1965.

Achterberg, C. van. 1976. A revision of the tribus Blancini (Hymenoptera,

Braconidae, Helconinae. Tijdschr. Ent. 118: 159-322.

Alford; Di. V.° 1975. ‘Bumblebees. 1-352.

Anonymous. 1963. Liste d'identification No. 5. Entomophaga 8: 333-373.

Capek, M. 1969. An attempt at a natural classification of the family

Braconidae based on various unconventional characters. Proc. Ent. Soc.

Wash. 71: 304-312.

Capek, M. 1970. A new classification of the Braconidae (Hymenoptera)

based on the cephalic structures of the final instar larva and biological

evidence. Can. Ent. 102: 846-875.

Fulmek, L. 1968. Parasitinsekten der Insektengallen Europas. Beitr. Ent. 18:

719-952.

Gruhl, K. 1967. Blacus-Tanze. Ergebnisse und Studien (Braconidae, Hymenop-

tera). Mitt. Deutsch. Ent. Ges. 26: 13-14.

Haeselbarth, E. 1973. Die Blacus-Arten Europas und Zentral-Asiens.

(Hymenoptera, Braconidae). Verdff. Zool. Staatssamml. Munchen 16:

69-170.

Konig, R. 1964. Brackwespen-Tanzschwarme der Gattung Blacus-erstmals in

Deutschland festgestellt (Hymenoptera: Braconidae). Faun. Mitt.

Norddeutschland 2: 209-210.

124 Contrib. Amer. Ent. Inst., vol. 20, 1983

Konig, R. 1967. Ein Beitrag zur Kenntnis einiger Braconidenarten in Schleswig-

Holstein (Hymenoptera: Braconidae, Blacinae). Faun.-Okol. Mitt. 3:

112-121.

Lipkow, E. 1965. Braconiden als Schmarotzer von Staphyliniden. Entomophaga

10: 377-381.

Marsh, P. M. 1959. Family Braconidae. In: Krombein, K. V. etal. Catalog

of Hymenoptera in America North of Mexico. Vol. 1: 144-295.

Mason, W. R. M. 1974. A generic synopsis of Brachistini (Hymenoptera:

Braconidae) and recognition of the name Charmon Haliday. Proc. Ent.

Soc. Wash. 76: 235-246.

Shenefelt, R. D. 1965. A contribution towards knowledge of the world litera-

ture regarding Braconidae (Hymenoptera). Beitr. Ent. 15: 243-500.

Shenefelt, R. D. 1969. Hymenopterorum Catalogus (nov. ed.) Part 4. Braconi-

dae |: 1-176. Junk, 's-Gravenhage.

Tobias, V. I. 1965. Genera groups and evolution of the subfamily Euphorinae

(Hymenoptera, Braconidae), Part 1. Entomol. Obozr. 44: 841-865 [in

Russian ].

KEY TO THE GENERA OF ORIENTAL MYMARIDAE, WITH A

PRELIMINARY CATALOG (HYMENOPTERA : CHALCIDOIDEA)

B. R. Subba Rao

Commonwealth Institute of Entomology

c/o British Museum (Natural History)

Cromwell Road, London SW7 S5BD, U.K.

M. Hayat

Department of Zoology, Aligarh Muslim University

Aligarh -— 202001, India

Abstract

The preliminary catalogue of Oriental Mymaridae, comprising India,

Pakistan, Sri Lanka, Nepal, Burma, Kampuchia, Malaysia, Indonesia and

Philippines is an up-to-date review. It includes 27 genera and 90 species.

All available types have been examined and this has necessitated many generic

transfers and specific synonymies. A key to genera with figures of species

representative of the genera dealt with in the catalog is presented.

INTRODUCTION

Species of the family Mymaridae are world-wide in their distribution and

some of these are the smallest known in nature. All the members of the

family so far known are parasitic in habit and develop in the eggs of other

insects, particularly of the Homoptera-~Heteroptera, but also of Lepidoptera,

Odonata, Coleoptera, Hemiptera and Psocids (Corrodentia). There are records

of mymarids having been reared from coccids and aleurodids, but these records

may in all probability involve misidentification of hosts. Mymarids have been

successfully utilized in the biological control of insect pests of crops and or-

chard trees and the outstanding example is the control of Gonipterus

scutellatus Gyllenhal in New Zealand and South Africa by the introduction

and establishment of Patassonnitens (Girault).

The mymarids not only are cosmopolitan in distribution but are also in

abundance. Yet these insects have not received the attention of entomologists

in general, and taxonomists in particular, possibly because of their minute size

and consequently not easily recognized and collected. Good tag mounts are

difficult to make of mymarids, and if made are hardly suitable for taxonomic

studies. It is necessary to prepare good micro slides for detailed studies.

Since Curtis described Mymar in 1832 over 140 genera with about 1250

species have been described and listed. However, only about 95 genera are

recognized. Mani (1938) in his catalog of Indian Chalcidoidea included only

5 genera and 5 species. Since then many genera and species have been de-

scribed and recorded by several Indian workers.

In this paper an effort has been made to bring together all the informa-

tion available on the Mymaridae of the Oriental Region comprising India,

Pakistan, Nepal, Burma, Sri Lanka and Malaysia. In compiling this catalog we

have been able to examine the types of several Oriental species available to

us through the courtesy of Dr. Grissell of the United States National Museum

and others in the British Museum (Natural History). These studies have neces-

sitated many generic transfers and some specific synonymies. However, in the

129

126 Contrib. Amer. Ent. dnst., vol. 20, 1983

case of large genera such as Camptoptera, Gonatocerus and Polynema, and

particularly those based only on males, it has not been possible to critically

examine their specific validity. The catalog includes 27 genera and 90 species.

The fauna being very rich in this region, there is no doubt that many more

genera and species known will be brought to light in addition to new forms

to science that will have to be described. It is hoped that the key to genera

of the Oriental Region and the numerous figures of species appended will en-

able workers of the Region to recognize and classify the mymarids they are

dealing with.

CLASSIFICATION

Mymarids were first treated as bona fide 'chalcids' as early as 1839 by

Haliday who also regarded them as a 'family', Mymaridae. Later authors,

such as Walker, Westwood and Foerster, regarded mymarids as Proctotrupoids.

This view was nearly abandoned since Ashmead (1904) has shown their chalcid

affinities.

Ashmead (1904) divided Mymaridae into two subfamilies, the pentamerous

Gonatocerinae and the tetramerous Mymarinae, and each of these subfamilies

into two tribes based upon the nature (mode) of attachment of the gaster with

the propodeum. Tribes Gonatocerini (gaster sessile or subsessile) and Ooctonini

(gaster petiolate) in Gonatocerinae, and Mymarini (gaster petiolate) and

_ Anaphini (gaster sessile or subsessile) in Mymarinae. Girault (1911, 1912) in

his earlier contributions on the North American and Australian mymarid fauna,

followed Ashmead's classification, but in his privately printed paper (1929)

dealing with the North American mymarids, considered the mode of attachment

of the gaster with the propodeum more important and relegated the tarsal

segments to tribal status. Thus he divided mymarids into two subfamilies:

Mymarinae with convexly rounded, subsessile or petiolate gaster, and Alaptinae

with truncately sessile gaster. Each one ot these two subfamilies was further

divided into two tribes based on the number ot tarsal segments; tribes

Mymarini (tetramerous) and Ooctonini (pentamerous) in Mymarinae, and Alap-

tini (pentamerous) and Anagrini (tetramerous) in Alaptinae.

Ghesquiére (1942) considered mymarids as an intermediate group between

Chalcidoidea and Proctotrupoidea, gave them the status of a super-family,

Mymaroidea, and recognized three families, Mymaridae, Lymaenonidae and

Signiphoridae.

Debauche (1948) placed Mymaridae in Chalcidoidea, and recognized

Lymaenonini and Mymarini as tribes of the family. He also proposed the

family Mymarommidae for the genus Mymaromma Girault (= Palaeomymar

(Meunier) which Girault (1920) did not assign to any tribe.

Annecke and Doutt (1961) followed Girault (1929) except they included

one more tribe, Anaphini, in the tetramerous Mymarinae. Yoshimoto, Kozlov

and Trjapitzin (1972) proposed a third subfamily, Eubroncinae, for two genera,

Eubroncus and Stemmarotrum, both characterized by peculiarly shaped head and

enormously developed mandible. Nikol'skaja (1978) and Kozlov and Rasnitsyn

(1979) considered Palaeomymar as belonging to the proctotrupoid family

Serphitidae, the latter authors downgraded Mymarommidae (they spelled it as

Mymarommatidae) to a subtamily Mymarommatinae in Serphitidae. They

Subba Rao & Hayat: Oriental Mymaridae 127

further tried to show that mymarids are related more closely to Procto-

trupoidea than to Chalcidoidea.

In the present work, we have followed the classification of Girault (1929)

although this system and that of Ashmead (1904) may not reflect natural af-

finities of the included genera. For this same reason we do not consider

Anaphini sensu Annecke and Doutt (1961) a useful category. The subfamily

Eubroncinae is also not accepted here since we consider the genera included

in that subfamily as nothing more than anomalous members of Mymarinae.

We, however, note that further detailed studies on the genera are needed to

establish natural affinities and for a better system of classification.

The following abbreviations are used for the depositories given in the

catalog:

BMNH - British Museum (Natural History), London, England.

BPBM - Bernice P. Bishop Museum, Honolulu, Hawaii, U-S.A.

ECO ~- Entomology Collection, Ministry of Food and Agriculture, Ottawa,

Canada.

EUM~ - Entomological Laboratory, Faculty of Agriculture, Ehime University,

Matsuyama, Japan.

IARI - Division of Entomology, Indian Agricultural Research Institute, New

Delhi, India.

IEUN- - Istituto di Entomologia Agraria dell'Universita di Napoli, Portici,

Italy.

HSPA_ ~- Division of Entomology, Hawaiian Sugar Planters' Association,

Honolulu, Hawaii, U.S.A.

MSNG - Museo di Storia Naturale, Geneve, Italy.

MUN) - Entomology Laboratory, Faculty of Agriculture, Meijo University,

Nagoya, Japan.

QMB-~ - Queensland Museum, Brisbane, Australia.

RNSS_ - Royal Norwegian Society Science Letters, Museum Tronheim.

USNM - United States National Museum of Natural History, Washington D.C.,

S.A.

pas | -~ Zoological Survey of India, Calcutta, India.

Family MYMARIDAE Haliday

-Mymares (Tribus) Haliday, 1833, Ent. Mag. 1: 341.

Mymaridae (Family) Haliday, 1839, Hym. Synop. p. Il.- Walker, 1846, Ann. Mag.

Nat. Hist. 18: 49.- Ashmead, 1897, Proc. Ent. Soc. Wash. 4: 236.-

Ashmead, 1904, Mem. Carnegie Mus. |: 362.- Mani, 1938, Cat. Indian

Ins. 23: 141.- Debauche, 1948, Mem. Mus. Hist. Nat. Belg. 108: 7.- Annecke

& Doutt, 1961, Ent. Rev. Dept. Agric. Tech. Serv. Repb. S. Afr. No. 5:

2.- Peck, 1963, Can. Ent. Suppl. 30: 18.- Peck, Boucek & Hoffer, 1964,

Mem. Ent. Soc. Canada No. 34: 112.- Graham, 1969, Bull. Ent. Mus. (Nat.

Hist.) (Ent.) Suppl. No. 16: 12.- De Santis, 1979, Catalogo De Los Himen-

opteros Calcidoideos De America Al Sur De Los Estados Unidos: 360.-

Burks, In: Krombein et al., 1979, Catalog of Hymenoptera in North

America North of Mexico, Vol. 1: 1022.

128 Contrib. Amer. Ent. Inst., vol. 20, 1983

Mymarides (subfamily) Westwood, 1840, Intr. Modern Class. Ins. Vol. II, Synop.

p. 78 (in family Proctotrupidae).

Mymaroidae Foerster, 1856, Hym. Stud. Heft. II: 20, 27, 116.

Mymaroidea (Superfamily) Ghesquiere, 1942, Rev. Zool.-Bot. Afr. 36: 319 (part).

Lymaenonidae Ghesquiere, 1942, Rev. Zool.-Bot. Afr. 36: 319.

Mymarommidae Debauche, 1948, Mem. Mus. Hist. Nat. Belg. 108: 7.

Mymarinae Schmiedeknecht,-1909, Gen. Ins. 97: 5, 488.

Gonatoceridae Mani & Saraswat, 1973, Mem. Sch. Ent. St. Johns' Coll. Agra

NO. 2728,

KEY TO HIGHER CATEGORIES AND ORIENTAL GENERA OF MYMARIDAE

1. Gaster broadly attached with the propodeum; mesopostphragma plainly

Precis to. te acre UP ies) BZ) a ee ea ee ae :

pre Se bea a ae men Uae Ae le tue Si Subfamily Alaptinae.... 2

Gaster convexly rounded at base, with a more or less distinct petiole;

mesopostphragma not or hardly projecting into the gaster (Figs. 3, 4)

ON EN Ay aces Pome ea gh nl abe gu aN Subfamily Mymarinae... . 3

Ae. ete Se oseemmented: sen i eG. ey) HIDE AMS Ne OR lee 4

eter Sseearrented: Gc ee Trips Alaprint ws Sk.

Be hte EOP NEI ON eae ee Wwe eke heibe MYMArIAl. . cs 62% 13

Tee ORION COG a a LEE COO ONIN oe oo eee 8

4. Antennal clava solid (Fig. 6); scutellum followed by a pair of distinctly

separated plates (= postscutellum) (Fig. 8); male antenna 13-segmented

PERT A Us a at A PNR OORT Cs Luar tT hoe Anagrus Haliday

Antennal clava 2- or 3-segmented; postscutellum composed of a single

Pye ERR SR Sr Deere | rng oe et INS mags) be 5

5. Clava 2-segmented (Fig. 10); body dorsoventrally flattened .........

BY Re a ie GHC ECE Uta Werte tou Qe a alt gt iat ap Wome Platypatasson Ogloblin

6. Clava with sutures strongly oblique (Fig. 13); body not flattened .....

SS RUE ET CTR AEC A CRSP NR. GRR Gap t RGN: WER: lenyeN ry Sse kamu Stethynium Enock

Clava with sutures perpendicular; body strongly flattened ..........

BERLE NUDE are eae Hugi sare SBN e ga Te RL eo Raa MERI gt ge Platystethynium Oglobin

7. Funicle 5-segmented (Fig. 15); scutellum and postscutellum not strongly

sculptured; marginal vein short; hind margin of fore wing excised be-

neath the venation (Fig. 17); male antenna 10-segmented (Fig. 16) . .

ws PRR Re ae, Rite GaSe Sala EL ak PUNE Grea Sige ck SRR SSRI UR FN Alaptus Westwood

Funicle 6-segmented (Fig. 19); scutellum and postscutellum strongly

sculptured (Fig. 21); marginal vein long; hind margin of fore wing nor-

mally rounded (Fig. 20); male unknown......... Litus Haliday

Oe CIe SROTE. WlCeet Tham (OMP Wee ue a, die enue Ske i ge uC a a Gg 9

Petiole at least as long as wide (Fig. 22)

10.

ip.

12.

14.

by.

16.

18.

Subba Rao & Hayat: Oriental Mymaridae 129

Funicle 5-segmented, clava solid or apparently 2~ or 3-segmented (Fig.

23); marginal vein greatly elongate, the venation extending to half or

more the length,,of wing (Vig. 24.060. eax go yexses: Arescon Walker

Funicle 8-segmented, clava solid (Fig. 25); marginal vein not greatly

elongate and venation not extending beyond basal third of wing (Fig.

Boy WER ee ta hae ear erry ane Oe Sains Eno re te Gonatocerus Nees

Funicle 8-segmented, antenna ll-segmented; propodeum with well-developed

cermpe (Eee 2 ioe winnel upd oo geginm enue. . Ooctonus Haliday

Funicle 7-segmented; antenna 10-segmented; propodeum at most very

WOSMIY GONMOle a. Gua hac miei. kM. Olen BGA Wires... 11

Second segment of funical normally developed, not anelliform .......

bs he” oh a. go ta ca eb Gy pw OG aie cine alist TE Ble in Paral e Eomymar Perkins

Second segment of funicle, in both sexes, anelliform, sometimes so small

as to. bavoverionked (Mige: 40 A hie hi Aes) cae AP te, SO 12

Fore wing narrow and distinctly curved on both margins; stigmal vein

short, less than half the length of marginal and with a truncate apex

hod. Mabe FE. 1 aM Nhat Wie Mtacke ML Rte: SA ANCL AMA dou hata Camptoptera Foerster

Fore wing comparatively broad, not at all curved; stigmal vein long, more

than half the length of the marginal and narrowed at apex (Male only,

female unknowih)idy enipien. Gab caw Aoi Camptopteroides Viggiani

Gaster subpetiolate or subsessile, or petiole distinctly wider than long

CPG es MS MR ety i TIA ey aOR eaten Cars Ace TA oan 14

Gaster distinctly petiolate, the petiole at least as long as wide, usually

sit ch Ma Ol ERE RIM taslg eb idcholbi a ap oahu main tichla ey Mb inehe hia rdl Mace ubain Nei, 19

Funicle 8-segmented, clava solid; propodeum and first segment of gaster

with prodigiously developed carinae; each side of propodeum with a

Prodigiousiy branched sete 600 0 0 2 ee Ptilomyar Annecke & Doutt

Funicle 5- or 6-segmented; propodeum and first segment of gaster with-

out such carinae and propodeum without such setae ......... 15

Funicle 5-segmented, antenna 8-segmented (Fig. 35); hypopygium promi-

nent, extending to apex of gaster; male antenna |2-segmented . . .

Moe Tem Ope tee Oh OMe |G Oe ere ee Paralleloptera Enock

Funicle 6-segmented, antenna 9- or 10-segmented (Figs. 38, 42); other

characters seme ee differant’ sie is). poi taed bles, eo ie ae 16

Clava 2-segmented (Fig. 38); male antenna i2-segmented (Fig. 39)...

PEI a Sic a Sa arma Panna d) smal hy atin 1 yal MiaR Py VR yd alg ON he Zc Patasson Walker

AANA ROMs i as! gi gata ieee dame ausia. nia ca 4 Niel | Deeneanaaetel 17

Hypopygium not prominent, not extending to apex of gaster; fore wing

distinctly curved on both margins (as in Camptoptera, Figs. 32-34);

male antentia l2-seomented . 9. 000 Eofoersteria Mathot

Hypopygium prominent, extending to apex of gaster (Fig. 43); fore wing

broad, not curved; male antenna ds~segmented: (wisi. Git Le e-, 18

Hind wing very broad (Fig. 46); propodeum elongate; at least as long as

postscutellum; first segment of gaster strongly carinate on sides (Fig.

48); wings densely setose; metanotum not raised above level of

PISO a ah Nh Mae aa ee ae Erythmelus Girault

130

20.

21.

223

24.

26.

Contnb. Amer. Ent. inst.,- vol. 20, 1983

Hind wing narrow; propodeum shorter than postscutellum; first segment

of gaster not carinate; metanotum raised above level of propodeum

(Fig. 43); wings sparsely setose to subnudus . . .. Erythmelus Enock

Mandible long, as long as the length of face; head, in profile, triangular,

with the frontovertex and face forming an acute angle; the space be-

tween antennal sockets with a subrectangular shelf-like structure . .

oe ORI Gg a Eubroncus Yoshimoto, Kozlov & Trjapitzin

Mandible short, distinctly shorter than length of face; head, in profile,

not acutely triangular, the frontovertex convex or flat, more or less

convexly merging with face; space between antennal sockets without

ee a a cet ae aa aca 20

Hind wing abbreviated, filiform, or poorly developed, with only a few mar-

ginal cilia (Figs. 51-53); fore wing oar-shaped and distinctly petiolate

(Fig. 50); scape long and medially constricted (Fig. 49) ........

ee ge ee oe Sah ree ee i aces tes RUA ene Oe 2 Maman Carte

Hind wing neither abbreviated nor filiform; fore wing differently shaped,

HOt istic Devigidte fier OE eoL Hees OR ee adie Bee ow ok tek a 21

Hind leg with very long spine-like setae; hind coxa longer than petiole

or mid femur (Fig. 55); fore wing with the discal setae arranged in

curved and alternating strong and weaker rows (Fig. 56); last segment

of funicle like a segment of clava (Fig. 54). . Narayanella Subba Rao

Hind leg without such long setae; hind coxa shorter than petiole or mid

femur (Fig. 4); fore wing with the discal setae not arranged in curved

rows (Figs. 58, 59); clava clearly differentiated from last funicle seg-

Ra ee hc Re ea eee ess Gar eet eA, Gow a Se. Gi Sa 22

Pro-— and mesothoracic dorsum with gross, spine-like, blunt or indented

ee te I a he ee a Ae es PENS TAN RP a RE gene RS 23

Thoracic dorsum. With. short, -NOLimal. SETAE «oo ceils co cia unite) ome) aa ae aces Me 25

Propodeum with a mid-longitudinal furrow or canal .............

PIQponecum Witout SHC. Cala, OF TUITOWs.) yg co 4cci toned Meee vata mance’ vs 24

Propodeum with two keels converging above the petiolar insertion to form

a more or less distinct process (Fig. 4); discal setae of fore wing

often with enlarged bases (tormae); fore wing often banded (Figs. 58,

BO ee gee pliet dale deal a Bails Nita iol ag: lg losatine at Giclee Acmopolynema Ogloblin

Propodeum without keels; fore wing not banded. . .Chaetomymar Ogloblin

Propodeum with two keels converging above the petiolar insertion to form

a more or less distinct process; discal setae of fore wing often with

enlarged bases; fore wing often banded..... Acmopolynema Ogloblin

Propodeum with the keels absent or different; wing usually hyaline and

UR AMC ee i ena Mla ae niin alle he 26

Scape with distinct scale-like structures; marginal vein thin; prothoracic

spiracles placed mesad of normal position, on the line between pro-

POtiah Bic: MESOSCULUMc: cob ss ew cl ee kk ees Stephanodes Enock

Scape without scale-like structures; marginal vein thickened; prothoracic

AGE TROT TE UC baer elk Cale a sti oy aks eo ae oe Gin ba a mec lan se 27

Subba Rao & Hayat: Oriental Mymaridae 131

27. Fore wing with two distinct rows of discal setae on basal half ......

imi li oe a or Re ane oc se, fa ahs Oa Grangeriella Soyka

Fore wing with discal setae not arranged in only two rows on basal half

Polynema Haliday

Dace ae to he See ey ie, ee So, Pee ee See at ee Ce ek ee 7 a. Ne a? 2 Lie ee) fe te ee A a eee

ACMOPOLYNEMA Ogloblin, 1946 *

Acmopolynema bagicha (Narayanan, Subba Rao & Kaur) comb. nov. - India:

Delhi, Maharashtra, Tamil Nadu.

Polynema bagicha Narayanan, Subba Rao & Kaur, 1960: 886, M, F. Delhi,

India (IARI).

Mymarilla deccana Mani & Saraswat, 1973: 109, F. Poona, India (USNM).

syn. nov.

Polynema indopeninsularis Mani & Saraswat, 1973: 119, M, F. Berijam,

Kodaikanal Hills, India (USNM), syn. nov.

Polynema deccana (Mani & Saraswat): Subba Rao, 1976: 89.

Acmopolynema incognita (Narayanan, Subba Rao & Kaur) comb. nov. - India:

Delhi.

Maidliella incognita Narayanan, Subba Rao & Kaur, 1960: 889, F. Delhi,

India (IARI).

Polynema (Maidliella) incognita (Narayanan, Subba Rao & Kaur):

Narayanan & Subba Rao, 1961: 667.

ALAPTUS Westwood, 1839

Synonyms: Parvulinus Mercet, 1912; Metalaptus Malenotti, 1917.

Alaptus delhiensis Mani - India: Delhi. | |

Alaptus delhiensis Mani, 1942: 160, F. New Delhi, India (IARI).

Alaptus magnanimus Annandale - India: West Bengal.

Alaptus magnanimus Annandale, 1909: 299, M. Calcutta, India (ZSI).

Alaptus ramakrishnai Mani - India: Tamil Nadu.

Alaptus ramakrishnai Mani, 1942, M=F. Coimbatore, India (IARI).

Host: (?) Mealybug on coconut.

ANAGROIDEA Girault, 1915

Anagroidea himalayana (Mani & Saraswat) | comb.nov.- India: Himachal

Pradesh.

Anaphes himalayanus Mani & Saraswat, 1973: 101, F. Kalotop, Dalhousie,

India (USNM).

numba) Leper cited se

Detailed citation to genera are not given here since most of these are avail-

able in Annecke & Doutt (1961) and for genera described after that date we

have given the references.

F = Female. M = Male.

132 Contrib. Amer. Ent. Inst., vol. 20, 1983

[Males described as belonging to this species by Mani & Saraswat (1973)

actually belong to Patasson.]

ANAGRUS Haliday, 1833

Synonyms: Pteratomus Packard, 1864; Packardiella Ashmead, 1904;

Paranagrus Perkins, 1905.

Anagrus columbi Perkins - India: Andhra Pradesh.

Anagrus columbi Perkins, 1905: 198, F. Ohio, U.S.A. (HSPA).

Host: Sogatella furcifera (Horwath).

Anagrus dalhousieanus Mani & Saraswat - India: Himachal Pradesh.

Anagrus dalhousieanus Mani & Saraswat, 1973: 104, F. Khajjar, Dalhousie,

India (USNM).

Anagrus empoascae Dozier - India: Delhi.

Anagrus empoascae Dozier, 1932: 86, F. Damien, Haiti (USNM). Subba Rao,

1966: 189, F, M. Delhi.

Host: Empoasca devastans Distant.

Anagrus flaveolus Waterhouse - India: Orissa, Andhra Pradesh. Pakistan.

Sri Lanka. West Malaysia, Philippines.

Anagrus flaveolus Waterhouse, 1913: 87, M, F. Trinidae (BMNH).

Host Nilaparvata lugens Stal.

Anagrus optabilis (Perkins) - Fiji. India: Andhra Pradesh, Orissa.

Paranagrus optabilis Perkins, 1905: 199, M. Queensland, Australia (HSPA),

also Fiji.

Hosts: Perkinsiella saccharicida Kirkaldy; Brown plant hopper.

Anagrus osborni (Fullaway) - Philippines: Los Banos.

Paranagrus osborni Fullaway, 1919: 53, F, M. Los Banos, Philippines (BPBM).

Host: Peregrinusmaidis (Ashmead).

Anagrus perforator (Perkins) - Fiji. India: Orissa. Philippines.

Paranagrus perforator Perkins, 1905: 199, F. Fiji (HSPA).

Host: Nilaparvata lugens St&l; Tarophagus proserpina (Kirkaldy).

ARESCON Walker, 1846

Synonyms: Panthus Walker, 1846; Leimacis Foerster, 1847; Limacis Foerster,

1856; Xenomymar Crawford, 1913; Neurotes Enock, 1914.

Arescon enocki (Subba Rao & Kaur) - India: Bihar, Delhi, Gujarat, Himachal

Pradesh, Karnataka, Maharashtra.

Neurotes enocki Subba Rao & Kaur, 1959: 235, F. New Delhi, India (IARI).

Arescon enocki (Subba Rao & Kaur): Subba Rao, 1966: 187, F, M.

distribution.

Subba Rao & Hayat: Oriental Mymaridae 133

Host: Jassid eggs.

CAMPTOPTERA Foerster, 1856

Synonyms: Stichothrix Foerster, 1856; Pteroclisis Foerster, 1856;

Macrocamptoptera Girault, 1910; Congolia Ghesquiere, 1942.

Camptoptera ambrae Viggiani - India: Tamil Nadu.

Camptoptera ambrae Viggiani, 1978b: 152, F. Kodaikanal, India (MSNG).

Camptoptera cloacae Taguchi - Philippines: Bohol Is.

Camptoptera cloacae Taguchi, 1972: 228, F. Longon, Bohol Is., Philippines

(MUN).

Camptoptera concava Taguchi - Philippines: Bohol Is.

Camptoptera concava Taguchi, 1972: 225, F. Bilar, Bohol Is., Philippines

(MUN).

Camptoptera longifuniculata Viggiani - India: Tamil Nadu.

Camptoptera longifuniculata Viggiani, 1978b: 153, F. Kodaikanal, India

(MSNG).

Camptoptera okadomei Taguchi - Philippines: Bohol Is.

Camptoptera okadomei Taguchi, 1972: 224, F. Longon, Bohol Is., Philippines

(MUN).

Camptoptera philippina Taguchi - Philippines: Bohol Is.

Camptoptera philippina Taguchi, 1972: 223, F. Longon, Bohol Is.,

Philippines (MUN).

Camptoptera protuberculata Viggiani - Sri Lanka:

Camptoptera protuberculata Viggiani, 1978b: 155, F. Mululla, Sri Lanka

(MSNG).

Camptoptera sakaii Taguchi - Taiwan.

Camptoptera sakaii Taguchi, 1977b: 143, F. Pinting Hsien, Taiwan (MUN).

Camptoptera serenellae Viggiani - Sri Lanka:

| Camptoptera serenellae Viggiani, 1978b: 151, M. Hakgala, Sri Lanka (MSNG).

Camptoptera taenia Taguchi - Philippines: Bohol Is.

Camptoptera taenia Taguchi, 1972: 225, M. Longon, Bohol Is., Philippines

(MUN).

Camptoptera taiwana Taguchi - Taiwan.

Camptoptera taiwana Taguchi, 1977b: 144, F. Pinting Hsien, Taiwan (EUM).

Camptoptera tuberculata Viggiani - Sri Lanka: |

Camptoptera tuberculata Viggiani, 1978b: 154, F. Ambagaswewa, Sri Lanka

(MSNG).

Camptoptera vinea Taguchi - Philippines: Bohol Is.

134 Contrib. Amer. Ent. Inst., vol. 20, 1983

Camptoptera vinea Taguchi, 1972: 227, M. Longon, Bohol Is., Philippines

(MUN).

CAMPTOPTEROIDES Viggiani, 1974

Camptopteroides armata Viggiani - Sri Lanka:

Camptopteroides armata Viggiani, 1974: 24, M. Hatton, Sri Lanka (MSNG).

CHAETOMYMAR Ogloblin, 1946

Chaetomymar elisabethae (Ferriere) comb. nov. - Java.

Polynema elisabethae Ferriere, 1931: 294, F. Buitenzorg, Java (BMNH).

Host: Euproctus flexuosa Snell.

Chaetomymar tayalum Taguchi - Taiwan.

Chaetomymar tayalum Taguchi, 1957b: 113, F. Fenchihu, Chiae Hsien, Taiwan

(EUM).

CLERUCHUS Enock, 1909

Sp. indet Malaysia, Johore.

Host: Callimerus: arufer.

EOFOERSTERIA Mathot, 1966

Eofoersteria secunda Viggiani - India: Tamil Nadu.

Eofoersteria secunda Viggianai, 1978a: 39, F. Kodaikanal, India (MSNG).

EOMYMAR Perkins, 1912

Eomymar muiri Perkins - Java.

Eomymar muiri Perkins, 1912: 27, F. Pekalongan, Java (BPBM).

Host: Delphacid, probably

ERYTHMELUS Enock, 1909

Synonym: Enaesius Enock, 1909.

Erythmelus empoascae Subba, Rao - India: Delhi.

Erythmelus empoascae Subba Rao, 1966: 192, M, F. Delhi, India (IARI).

Host: Empoasca devastans Distant.

Erythmelus helopeltidis Gahan - India: Andhra Pradesh, Uttar Pradesh. Malaysia:

Kuala Lumpur.

Erythmelus heopeltidis Gahan, 1949: 75, M, F. Kuala Lumpur, Malaysia

(USNM). Subba Rao, 1970: 662, F. India.

Hosts: Helopeltis cinchonae Mann., Ophiomyia lantanae.

EUBRONCUS Yoshimoto, Kozlov & Trjapitzin, 1972

Subba Rao & Hayat: Oriental Mymaridae 135

Eubroncus orientalis Yoshimoto, Kozlov & Trjapitzin — Malaysia: Selangor.

Eubroncus orientalis Yoshimoto, Kozlov & Trjapitzin, 1972: 880, M.

Selangor, Malaysia (ECO).

GONATOCERUS Nees, 1834

Synonyms: Lymaenon Walker, 1846;

Rachistus Foerster, 1847; Cosmocomoidea Howard, 1908;

Oophilus Enock, 1909; Agonatocerus Girault, 1913;

Gonatoceroides Girault, 1913; Gastrogonatocerus Ogloblin, 1935.

Gonatocerus bakrotus Mani & Saraswat - India: Himachal Pradesh.

Gonatocerus bakrotus Mani & Saraswat, 1973: 80, F. Dalhousie, India (USNM).

Gonatocerus berijamus Mani & Saraswat - India: Tamil Nadu.

Gonatocerus berijamus Mani & Saraswat, 1973: 82, M, F. Anamalai Hills,

India (USNM).

Gonatocerus bifasciativentris Girault - Java.

Gonatocerus bifasciativentris Girault, 1917: 134, M,F. Java (?USNM).

Host: ‘Eggs of a leaf-hopper embedded in the leaves of sugar-cane.'

Gonatocerus brevifuniculatus Subba Rao - Indonesia: East Sumatra. Pakistan:

Hyderabad.

Gonatocerus brevifuniculatus Subba Rao, 1970: 664, M, F. Hyderabad,

Pakistan (BMNH), also East Sumatra.

Host: ?Scale on Albizia sp.

Gonatocerus delhiensis (Narayanan & Subba Rao ) comb. nov. - India: Delhi.

Lymaenon delhiensis Narayanan & Subba Rao, 196]: 659, F. Delhi, India

(IARI).

Gonatocerus devikulamus Mani & Saraswat - India: Tamil Nadu.

Gonatocerus devikulamus Mani & Saraswat, 1973: 84, F. Berijam, Anamalai

Hills, India (USNM).

Gonatocerus devitatakus Mani & Saraswat - India: Tamil Nadu.

Gonatocerus devitatakus Mani & Saraswat, 1973: 86, F. Devikulum,

Anamalai Hills, India (USNM).

Gonatocerus empoascae (Subba Rao) comb. nov. - India: Pubjab.

Lymaenon empoascae Subba Rao, 1966: 195, F. Jullunder, India (IARI).

Host: Empoasca devastans Distant.

Gonatocerus indicus (Subba Rao & Kaur) comb. nov. - India: Delhi.

Lymaenon indicus Subba Rao & Kaur, 1959: 229, M, F. New Delhi, India

(IARI). :

136 Contrib. Amer. Ent. Inst., vol. 20, 1983

Gonatocerus kanheriensis Mani & Saraswat ~— India: Maharashtra.

Gonatocerus kanheriensis Mani & Saraswat, 1973: 87, F. Bombay, India ©

(USNM).

Gonatocerus longicrus Kieffer - India: Orissa.

*Gonatocerus longicrus Kieffer, 1913: 201, M.

Gonatocerus lucidus Dodd - Java.

Gonatocerus lucidus Dodd, 1919: 161, M, F. Tjibodas, Java (BMNH).

Gonatocerus munnarus Mani & Saraswat - India: Tamil Nadu.

Gonatocerus munnarus Mani & Saraswat, 1973: 89, F. Munnar, Anamalai

Hills, India (USNM).

Gonatocerus narayani (Subba Rao & Kaur) comb. nov. - India: Delhi.

Lymaenon narayani Subba Rao & Kaur, 1959: 227, F. Delhi, India (IARI).

Gonatocerus nigriodes (Narayanan & Subba Rao) comb. nov. - India: Delhi.

Lymaenon nigrioides Narayanan & Subba Rao, 1961: 656, F. Delhi,India

(IARI).

Gonatocerus pahlgamensis (Narayanan) comb. nov. - India: Jammu & Kashmir.

Lymaenon pahlgamensis Narayanan, 1961: 25, F. Kashmir, India 9IARI).

Host; ?Quadraspidiotus perniciosus (Comstock).

Gonatocerus ramakrishnai (Subba Rao & Kaur) comb. nov. - India: Delhi.

Lymaenon ramakrishnai Subba Rao & Kaur, 1959: 232, M, F. Delhi, India

(IARI).

Gonatocerus relictus Mani & Saraswat —- India: Tamil Nadu.

Gonatocerus relictus Mani & Saraswat, 1973: 90, M, F. Berijam, Anamalai

Hills, India (USNM).

Gonatocerus sahadevani (Subba Rao & Kaur) comb. nov. - India: Delhi.

Lymaenon sahadevani Subba Rao & Kaur, 1959: 231, F. Delhi, India (IARI).

Gonatocerus sarawakensis Sveum, Malaysia, Borner, Sarawak.

Gonatocerus sarawakensis Sveum, 1981: 81, M, F. Semengo-Soult & Kuching

(RNSS).

Gonatocerus shasthryi (Subba Rao & Kaur) comb. nov. - India: Karnataka.

Lymaenon shasthryi Subba Rao & Kaur, 1959: 234, M, F. Mandya, India

(IARI).

Host; Tettigonella spectra (Distant).

Gonatocerus tamilanus Mani & Saraswat - India: Tamil Nadu.

Gonatocerus tamilanus Mani & Saraswat, 1973: 93, M, F. Madras, India

(USNM).

“Type depository not traced.

Subba Rao & Hayat: Oriental Mymaridae tof

Gonatocerus tarae (Narayanan & Subba Rao) comb. nov. - India: Delhi.

Lymaenon tarae Narayanan & Subba Rao, 1961: 657, F. Delhi, India (IARI).

Gonatocerus udakamandus Mani & Saraswat - India: Tamil Nadu. ;

Gonatocerus udakamandus Mani & Saraswat, 1973: 96, F. Ootacamund, India

(USNM).

Gonatocerus uttarodeccanus Mani & Saraswat - India: Tamil Nadu.

Gonatocerus uttarodeccanus Mani & Saraswat, 1973: 97, M, F. Munnar,

Palghat, India (USNM).

GRANGERIELLA Soyka, 1956

Grangeriella indochinensis Soyka ~ Campuchia (Saigon).

Grangeriella indochinensis Soyka, 1956: 18, F. Saigon, Ho Chi Min

(Soyka's coll.)

HIMOPOLYNEMA Taguchi, 1977

Himopolynema malayanum Taguchi - Malaysia.

Himopolynema malayanum Taguchi, 1977a: 139, M. Tanah Rata, West Malaysia

(MUN).

Himopolynema parviscutum Taguchi - Taiwan.

Himopolynema parviscutum Taguchi, 1977a: 140, M. Roushing Spa, Taiwan

(EUM).

Himopolynema taiwanum Taguchi - Taiwan.

Himopolynema taiwanum Taguchi, 1977a: 142, M. Kenting Park, Taiwan (EUM).

LITUS Haliday, 1833

Litus enocki Howard - Sri Lanka.

Litus enocki Howard, 1896: 643, F. Pundauloya, Sri Lanka (USNM).

Host: ?Eriococcus sp.

MYMAR Curtis, 1832

Synonyms: Pterolinononyktera Malac, 1943; Oglobliniella Soyka, 1946.

Mymar schwanni Girault - India: Uttar Pradesh.

Mymar schwanni Girault, 1912: 166, F. Victoria, Australia (QMB). Verma,

1980: 536, F. Aligarh.

Mymar taprobanicum Ward - Widely distributed throughout the Oriental Region.

Mymar taprobanicum Ward, 1875: 197, F. Sri Lanka * (Type locality?).

Taguchi, 1975a: 22, Taiwan. Subba Rao, 1976: 89, synonymy. Hayat,

1977: 283, F, Jodhpur.

agrees eed

Type depository not traced.

138 Contrib. Amer. Ent. Inst., vol. 20, 1983

Mymarilla taprobanicum (Ward): Mani, 1938: 143.

Mymar indica Mani, 1942: 160, M. Delhi, India (IARI). Mani & Saraswat,

1973: 107, M. Coimbatore.

Host: Asociated with thrips. [In Japan one female bred from

Fallen. ] |

NARAYANELLA Subba Rao, 1976

Synonym: Narayana Subba Rao, 1976; not Narayana Distant, 1908.

Narayanella pilipes (Subba Rao) - Burma.

Narayana pilipes Subba Rao, 1976a: 88, M, F. Burma (BMNH).

Narayanella pilipes (Subba Rao): Subba Rao, 1976b: 352.

Narayanella thornypoda (Narayanan & Subba Rao) - India: Bihar, Karnataka,

Maharashtra.

Polynema (Polynema) thornypoda Narayanan & vote Rao, 1961: 665, M.

Balehonnur, India (IARI).

Anagrus khandalus Mani & Saraswat, 1973: 105, F. Bombay, India (USNM).

Saraswat & Mukerjee, 1975: 62, F. Ranchi.

Narayana thornypoda (Narayanan & Subba Rao): Subba Rao, 1976a: 89,

synonymy.

Narayanella thornypoda (Narayanan & Subba Rao): Subba Rao, 1976b: 352.

OOCTONUS Haliday, 1833

Synonym: Sphecomicrus Walker, 1846.

Ooctonus kodainus Mani & Saraswat - India: Tamil Nadu.

Ooctonus kodainus Mani & Saraswat, 1973: 78, F. Anamalai, India

(USNM).

PARALLELAPTERA Enock, 1909

Synonym: Anthemiella Girault, 19IlI.

Sp. indet: India, Karnataka, Bangalore.

Host: Dictyla sp. Teleonemia’scrupulosa ion Lanatana camera.

PATASSON Walker, 1846

Synonyms: Anaphoidea Girault, 1915; Anaphoides Enock, 1915; Yungaburra

Girault, 1915; Hofenderia Soyka, 1949; Fulmekiella Soyka, 1949;

Schizophragma Ogloblin, 1949.

Sp. indet, India: Uttar Pradesh, Himachal Pradesh.

PLATYPATASSON Ogloblin, 1946

Platypatasson fransseni Ogloblin - Indonesia (Java).

Platypatasson fransseni Ogloblin, 1946: 293, F. Java (USNM).

Subba Rao & Hayat: Oriental Mymaridae 3 139

PLATYSTETHENIUM Ogloblin, 1946

Platystethenium onomarchicidum Ogloblin - Indonesia (Java).

Platystethenium onomarchicidum Ogloblin, 1946: 291, F. Java (USNM).

POLYNEMA Haliday, 1833

Synonyms: Eutriche Nees, 1834; callitriche Agassiz, 1848; Cosmocoma

Foerster, 1856; Valkerella Westwood, 1878; Walkerella Westwood, 1883;

Maidliella Soyka, 1946; Novickyella Soyka, 1946.

Polynema ahlaensis Mani & Saraswat - India: Himachal Pradesh.

Polynema ahlaensis Mani & Saraswat, 1973: lll, F. Dalhousie, India (USNM).

Polynema anamalaiensis Mani & Saraswat - India: Tamil Nadu.

Polynema anamalaiensis Mani & Saraswat, 1973: 113, M. Palghat, India

(USNM).

Polynema anantanagana Narayanan - India: Jammu & Kashmir.

Polynema anantanagana Narayanan, 1961: 24, F. Anantnag, India (IARI).

Host: ? Quadraspidiotus perniciosus (Comstock).

Polynema crassa Mani & Saraswat - India: Himachal Pradesh.

Polynema crassa Mani & Saraswat, 1973: 114, M. Dalhousie, India (USNM).

Polynema dhenkunda Mani & Saraswat - India: Himachal Pradesh.

Polynema dhenkunda Mani & Saraswat, 1973: 116, M, F. Dhenkund, Dalhousie,

India (USNM).

Polynema indica Narayanan & Subba Rao - India: Delhi.

Polynema (Polynema) indica Narayanan & Subba Rao, 1961: 663, F. Delhi,

India (IARI).

Polynema kalatopensis Mani & Saraswat - India: Himachal Pradesh.

Polynema kalatopensis Mani & Saraswat, 1973: 122, M. Kalatop, Dalhousie,

India (USNM).

Polynema kamathi Mani & Saraswat - India: Himachal Pradesh.

Polynema kamathi Mani & Saraswat, 1973: 123, F. Dalhousie, India (USNM).

Polynema loriger Kieffer - Philippines.

Polynema loriger Kieffer, 1916: 290, M. Luzon, Philippines, type depository

not traced.

Polynema mendeli Girault - Queensland, Australia.

Polynema mendeli Girault, 1913: 118, M, 99 F. Nelson (Cairns) (QMB).

Polynema oophaga Subba Rao, 1970: 666, F, M. Malaysia (BMNH).

Host: Tettigoniid eggs on Paddy.

Polynema orientalis (Narayanan, Subba Rao & Kaur) - India: Delhi.

140 Contrib. Amer. Ent. Inst., vol. 20, 19838

Maidliella orientalis Narayanan, Subba Rao & Kaur, 1960: 888, M, F.

Delhi, India (IARI).

Polynema (Maidliella) orientalis (Narayanan, Subba Rao & Kaur):

Narayanan & Subba Rao, 1961: 667.

Polynema robusta Sveum - Malaysia, Indonesia.

Polynema robusta Sveum, 198]: 84, F. Pandu, Malaysia (RNSS).

Polynema truncata Narayanan & Subba Rao - India: Delhi.

Polynema (Polynema) truncata Narayanan & Subba Rao, 196]: 664, F. Delhi,

India (IARI).

PTILOMYMAR Annecke & Doutt, 1961

Ptilomymar besucheti Viggiani - Sri Lanka.

Ptilomymar besucheti Viggiani, 1974: 27, M, F. Mahaweli Ganga, near

Kandy, Sri Lanka (MSNG). |

Ptilomymar orientalis Taguchi — Philippines: Bohol Is.

Ptilomymar orientalis Taguchi, 1972: 229, M. Longon, Bohol Is.,

Philippines (MUN).

STEPHANODES Enock, 1909

Stephanodes imbricatus (Narayanan & Subba Rao) comb. nov. - India: Delhi.

Polynema (Stephanodes) imbricatus Narayanan & Subba Rao, 1961: 667, F.

Delhi, India (IARI).

Stephanodes orientalis Taguchi ~— Taiwan.

Stephanodes orientalis Taguchi, 1978: 73, M, F. Ishihara near Fukuoka,

Japan (EUM), also Taiwan.

STETHYNIUM Enock, 1909

Stethynium empoascae Subba Rao - India; Delhi.

Stethynium smpoascae Subba Rao, 1966: 189, M, F. Delhi, India (IARI).

Host: Empoasca devastans Distant.

ACKNOWLEDGMENTS

One of us (M. Hayat) is thankful to the Trustees of the British Museum

(Natural History), London, and Dr. L. A. Mound, Keeper of Entomology, BMNH,

for permission to study the Museum collections and for necessary facilities.

He is thankful to the Royal Society and the Commonwealth Foundation, London,

for financial assistance during the study period (July 1981 - June 1982) at the

BMNH.

Subba Rao & Hayat: Oriental Mymaridae 141

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144

ocetput

postertor ocellar ltne

antertor ocellar line

oceular line’

antemmal socket

° Co eS

medtan See

suborbttal carina

frontal carina

subocular Ai mandtble

suture 1 maxillary palp

pronotum —

notaular ltne

‘stde lobe (=parapsis)

mesoscutum

axtlla

scutellum

Tae senstlLla -

anterscutelLum

En BST epee

postscutellum

metanotum

propodeal sptracle

propodeum

pettole

-gaster

marginal vein

submarginal vetn

macrochaeta A

mtcrochaeta

sttgmal vein / hypochaeta

Figs. 1-5. 1, Generalized Mymarid, head in frontal view. 2, Gen-

eralized mymarid, head dorsum. 3, Gonatocerus sp., F, thoracic dorsum.

4, Acmopolynema bagicha, F, thoracic dorsum. 5, Generalized mymarid,

fore wing venation.

Subba Rao & Hayat: Oriental Mymaridae 145

Figs. 6-14. 6,7, Anagrus optabilis, F:

of fore wing.

6, antenna; 7, basal part

8, A. flaveolus, F, thoracic dorsum and base of gaster.

9-12, Platypatasson fransseni: 9, head dorsum, F; 10, antenna, F; 11, an-

tenna, M; 12, basal part of fore wing, F.

13,14, Stethynius triclavatum, F:

13, last segment of funicle and clava;

14, basal part of fore wing.

146 Contribs Amer. Ent.inst.,. vol. 20, 1983

sie SRT

Sewanee :

ens eke sats gc

3 Ss beers

EES Fi a - Ln Se: Were erates Jett

< et OE Se i A

—~—— =

HO ye 21 23

Figs. 15-23. 15-18, Alaptus antillantus: 15, antenna, F holotype;

16, antenna, M; 17, fore wing, F; 18, hind wing, F. 19-21. Litus cynifrons

F: 19, antenna; 20, basal part of fore wing; 21, part of thorax and gaster.

22, Ooctonus sublaevis, M, part of thorax and gaster. 23, Arescon enocki,

F, antenna.

Subba Rao & Hayat: Oriental Mymaridae 147

Figs. 24-34. 24, Arescon enocki, F, tore wing. 25, Gonatocerus sp.,

F, antenna. 26, G. shasthryi, F, pedicel and basal funicle segments. 20,

G. brevifuniculatus, F, holotype, antenna with distal funicle segments and

Clava omitted. 28-29, Gonatocerus sp.: 28, basal two flagellar segments,

M; 29, basal part of fore wing, F. 30-34, Camptoptera sp.: 30, antenna with

second funicle segment enlarged and shown separately, F; 31, antenna showing

ring-like second and fourth flagellar segments, M; 32, fore wing,F; 33, basal

part of fore wing, F; 34, hind wing, F-.

148 Contrib. Amer. Ent. Inst.,; vol. 20, 1983

hypopygtum

Figs. 35-43. 35-37, Parallelaptera sp., F: 35, antenna; 36, fore wing;

37, hind wing. 38-41, Patasson sp.: 38, antenna, F; 39, antenna, M; 40,

part of thorax and gaster showing the anteriorly extended inner valvulae;

41, basal part of fore wing, F. 42-43, Erythmelus helopeltidis, F : 42,

antenna; 43, part of thorax and gaster.

Subba Rao & Hayat: Oriental Mymaridae 149

Figs. 44-53. 44-48, Anagriodea sp.: 44, antenna, F; 45, antenna, M;

46, hind wing, F; 47, basal part of fore wing, F; 48, part of thorax and

gaster. 49-51, Mymar taprobanicum, F : 49, antenna; 50, fore wing; 51,

hind wing. 52, M. regalis, hind wing, F. 53, M. schwanni, hind wing, F.

150 —. ConteibArmmet..Emty drsts, vale 20, 1983

= POX

Figs. 54-62. 54-56, Narayanella sp., F: 54, antenna; 55, part of body;

56, fore wing. 57-58, Acmopolynema incognita, F: 57, antenna; 58, fore wing.

59, A. bagicha, F, fore wing. 60, Chaetomymar elisabethae, F holotype, basal

part of fore wing. 61-62, Polynema spp., basal part of fore wing in two

species, F.

THE BIOLOGY OF THE EUROPEAN SPECIES OF THE

GENUS ICHNEUMON AND RELATED SPECIES

(HYMENOPTERA : ICHNEUMONIDAE)

Rolf Hinz

Fritz—Reuter Str. 34, D-3352 Einbeck, BRD.

There are many incorrect statemients on the biology and hosts of the

Ichneumon species in literature. Misidentifications are common, both in

the names of the hosts and the ichneumonid species involved. The species

of Ichneumon emerge from the host pupae and the identification of the

pupae is not always easy.

To investigate the biology of the Ichneumon species, it became neces-

sary to conduct experiments on oviposition. This was facilitated by the fact

that nearly all Ichneumon species hibernate as gravid females. (Till now |

have found only one exception: Ichneumon deliratorius L. hibernates in

the host pupa). These hibernating females can easily be found from autumn

till spring in well known places (Oehlke, 1967: 77; Hinz, 1968: 77) and with

a suitable method it is possible to keep them in the refrigerator without

mortality. Thus the breeding of the Ichneumon species can be attempted.

It is possible without great difficulties if at the correct moment suitable

hosts are available in sufficient numbers and of the right age. Thus the

breeding of the Ichneumon species is reduced to a breeding of a large num-

ber of larvae and pupae of different Lepidoptera, which can be offered to

the parasite.

As far as known all Ichneumoninae are parasites of the Lepidoptera .

The assumption of Townes, that the oxypygous Ichneumoninae parasitize

pupae and that the hosts of the amblypygous species are the larvae has been

confirmed in all the cases.. Sometimes the oxypygous species prick the pre-

pupa (e.g. Ichneumon caloscelis Wesm. of several Satyridae). They move

back from the intensely moving caterpillar and do not parasitize them perhaps

because of the different structure of the surface. Some species accept, or

perhaps prefer, recently transformed, often still uncolored pupae (e.g.

Ichneumon nebulosae Heinz on Polia nebulosa Hufn.: Heinz, 1975: 256).

This seems to be astonishing for the pupae lying in holes in the ground, but

it is easily understood by the parasite's method to find the host: It fol-

lows the smell-trail the caterpillar leaves when it hides. Since the smell is

not perceptible for a long time, there is a great probability for the parasite

to find a young pupa.

When the parasite has entered the hole or found the cocoon of the

host, the abdomen is bent and the ovipositor, supported by the ovipositor

sheaths, is inserted vertically on the surface of the pupa. In fresh or

slightly sclerotized pupae the pressure of the ovipositor is sufficient. Species

that prick older or heavily sclerotized pupae, often have difficulties in ovi-

position. Turning the ovipositor back and forth helps its insertion into the

pupa, when the female presses its back against the side of the cocoon. As

a rule, after oviposition the female turns around and sucks the haemolymph

oozing out at the site of the prick.

151

152 Contrib. Amer. nt. Inst., vol. 20, 1983

Most of the European species have only one generation per year. That

causes an unusual long duration of life of more than eleven months for the

female, since the development from oviposition till hatching of the imago re-

quires in species of medium size around 25 days. During this time the fe-

males must feed, especially nectar of flowers, like those of the Umbelliferae.

At suitable times probably all species can be found in such places.

Some of the common species, like Ichneumon confusor Grav., melanotis

Hlgr., molitorius L., etc. have at least the potentiality of having several

generations per year. This is regulated by photoperiodism: Under long day

condition (16 hours) the ovaries of the females are fully developed 8-l4 days

after hatching. These species necessarily seem to be polyphagous, since

univoltine species often are monophagous or oligophagous. It is difficult to

state how many generations really appear in nature, since this can only be

concluded from the appearance of fresh males. Ichneumon males can hardly

be determined to species.

The ovaries of the hibernating females develop in spring with the in-

crease of temperature. There is a real diapause during the winter, since it

is rarely possible to induce the growth of the ovaries earlier by rise of

temperature. It is probable that all Ichneumon species have arrhenotok

parthenogenesis: in all species that I tested in this respect, the eggs of un-

fertilized females gave rise to males only.

Most of the species search for their hosts soon after hibernation in

spring. They prefer the pupae of Lepidoptera hibernating as larvae (i.e.

species of the genus Amathes and related species) or of butterflies flying

early in the year (e.g. Ichneumon inquinatus Wesm. as parasite of several

species of the genus Orthosia). Perhaps exceptions may be found in this

respect. At this time I only know one in a genus related to Ichneumon :

Chasmias paludator Desv. hibernates as imago, but oviposits not before

August. The female parasitizes the pupae of butterflies of the genus

Nonagria and Archanara living in the stems of Typha or Phragmites communis.

It opens with its jaws the "windows" the caterpillar has prepared and thus

reaches the pupa.

REFERENCES

Hinz, R. 1968. Die Untersuchung der Lebensweise der Ichneumoniden mit

Anhang: Bemerkungen zur Praeparation von Ichneumoniden. Ent. Nachr.

12: 73-81.

Hinz, R. 1975. Beschreibung und Zucht von Ichneumon nebulosae spec.

nov. Beitr. Ent. 25: 255-256.

Oehlke, J. 1967. Fang, Zucht und Praeparation von Schlupfwespen. Ent.

Nachr. li: 69-87.

A REVISION OF WESTERN PALEARCTIC OXYTORINE GENERA

PART III GENUS PROCLITUS

(HYMENOPTERA, ICHNEUMONIDAE)

G. van Rossem

Plant Protection Service, Wageningen, The Netherlands

Redescriptions of the Western Palearctic species of Proclitus Forster,

1868 are given together with a key to the species.

GENUS PROCLITUS

Cryptus (Clepticus) Haliday, 1838. Ann. Nat. Hist. 2: 112-121,

Proclitus Foerster, 1868. Verh. Naturh. Ver. Preuss. Rheinl. 25: 172.

Proclitus: Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl. 28: 113.

Plectiscus (Proclitus): Thomson, 1888. Opusc. Ent. 37 (Fasc. 12): 1306-1307,

Proclitus: Townes, 1971. The Genera of Ichneumonidae Part 4: 194,

Proclitus: Aubert, 1977. Spixiana 1(2): 142.

KEY TO THE MALES

The males of P. fulvipectus and P. rudis are unknown and are not in-

cluded in the key. I have not seen the males of P. fulvicornis and

comes, but the differences in the key rest on the females. The distinction

of P. attentus, P. praetorand P. albidipes is rather complicated and not

quite reliable.

Anterior tentorial pits impressed and with a carina between the eye margin

and. the Javetal, Commer at Wie hem i i ee peek ah ia Mee Mele uk Le

Anterior tentorial pits not impressed and with no carina between the eye

WATE, ONG INS Ci ee sie eS a ae a ee ue a 4

Clypeus flat, the apical margin arcuate, 2.5 x as wide as long... .....

to aay ope Ma: Bee acs ana dea ile os eugene Vig) He MaMa dak a Meee ne te, ee cee 4 . P. paganus

Wlypeus. convex, the apion) margin almost, atti Ntig « focciciaccic otc se ae. 3

Clypeus medially protruding, 1.8-2.4 x as wide as long ............

Sa ae ae We tar ge ig ae ary Ae Ter Mae Meas Bee URC ain PORTA YO a) OBL. falvioornis

Clypeus medially protruding, 2.3 x as wide as long. Carina between eye

Mareiy Bhd Chyene Ween sk a Ce our et eee is 2. Fe Comes

Face wide, about 0.45 of frontal width of head (including the eyes), with

a row of conspicuous long, erect setae along the inner margin of eye.

Clypeus exceptionally wide, almost 3 x as wide as long, the apical mar-

gin arcuate. Lateral corner of clypeus reaching beyond line of inner

margin of eye. Clypeus with widely placed long setae ..........

First tergite coriaceous. Lateral carinae of scutellum running to apex of

BOUL GLIUIN, TU I re ow kar fairent Pere ts GRO Sie PF... 2onatus

First tergite not coriaceous. Lateral carinae of scutellum not running to

apex of scutellum, only present at proximal corners of scutellum... 6

153

154 Gontrib. “Amer. Ent. inst., vol. 20, 1983

6. Length of postannellus : length of second flagellar segment = about 11 : 10

(postannellus slightly longer). First gastral segment 3.4 - 3.8 x as long

ASGABPICAMY WMG): Go 6 bh ek ee = 7 soP ~ attentus

Length of postannellus : length of second flagellar segment = 4: 3; 16: 13

OG 1S a ee ak a oo ee 7

. Length of postannellus : length of second flagellar segment = 4: 3 to 16:

13 (postannellus longer). First gastral segment 4.4 - 5.0 x as long as

apically wide . ar... e- SO a er et eer Lu. Pegpraetor

Length of postannellus : length of second flagellar segment = 13 : 10. In-

ner margins of eyes somewhat converging, width at antennal sockets :

Width at clypeus = cit 2 ge Se Se ee P. albidipes

KEY TO THE FEMALES

Ovipositor fone, 0.5 = > of leneih front’ wing... . .'sos ee ee 2

Ovipositor short, length below 0.6 of length front wing **......... 3

Ovipositor exceptionally long, 0.80 - 1.05 of length front wing. First gastral

segment about 2.5 as long as wide apically. Anterior tentorial pits im-

pressed, with a groove between eye margin and clypeus. Clypeus weakly

Convex, apour 2 4°x% as Wide as lone = oS ee 2. P. comes

Ovipositor not exceptionally long, 0.5 - 0.7 of length front wing. First

gastral segment very slender, about 4.3 x as long as wide apically. An-

terior tentorial pits somewhat impressed, with a groove between eye

margin and clypeus. Clypeus weakly convex, 1.7 - 2.4 x as wide as

POMC ek ee 6 hl he oe UE es cae ee es 1 Po practor

Anterior tentorial pits impressed and with a carina between eye margin and

Pletal Geer Ol Uy oe es 4

Anterior tentorial pits not impressed and with a weak or no carina between

ee ate eet ee ee, ee 5

Clypeus convex, the apical margin straight, medially protruding, 1.8 x as

wide as long. Ovipositor 0.27 - 0.44 of length front wing ........

See Rs ce ret er aur cg i a ies yd ae Gee ea eae Pian Ck 3. Pe. Fulvicornis

Clypeus flat, the apical margin arcuate, 2.5 x as wide as long. Ovipositor

0.24 ~ 0,36 of length fvent wing % 408.) -.cibe 2rd. ele. 4 . P. paganus

Ovipositor very short, 0.26 of length front wing. Malar space with a groove.

Lateral carinae of scutellum running to apex of scutellum, not meeting.

Apex of scutellum striated or with somewhat rought sculpture. First, and

in some specimens also the second tergite coriaceous. .5 . P. zonatus

Ovipositor longer, 0.36 - 0.48 of length front wing. Lateral carinae of

scutellum only at proximal.corners: of scutellum’s. 0... Pop eolo. . . 6

Anterior tentorial pits conspicuous, open. A carina and laterally of the

carina a groove between eye margin and lateral corner of clypeus.

—

Character not reliable in comparison with P. praetor males.

In some specimens of P. fulvicornis the length of the ovipositor exceeds

0.6 the length of front wing. These specimens have a wider malar space

(0.31 - 0.35 of width face) than in P. praetor.

van Rossem: Proclitus (Ichneumonidae) 155

Clypeus medially somewhat convex, about as wide as width face .. .

sha AE SE ga a Tee eG Oe Ces es a Pa 6. P. fulvipectus

Anterior tentorial pits not conspicuous. A groove between eye margin and

clypeus, but no carina present. Clypeus not as wide as face ..... 7

7. Radius originating at distal 0.54 of stigma. First gastral segment slender,

aDOUL SX pe lone ds Wwe apically... oo oe ae en 7, B. attentus

Radius originating at about 0.50 of stigma. First gastral segment rather

wide and short, 24 % as long, as, wide apically. se. on e. ao. Po yudie

1. Proclitus praetor (Haliday in Curtis)

Cryptus (Clepticus) praetor Haliday in Curtis, 1838. Ann. Nat. Hist.

2) 1h, P.

Proclitus grandis Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 116.

Proclitus praetor: Fitton, 1976. Bull. Brit. Mus. Nat. Hist. (Ent.) 32

(8): 333.

Proclitus praetor: Aubert, 1977. Spixiana 1(2): 142.

Characteristics of the lectotype of cryptus (Clepticus) praetor: Labels:

lectotype label M. G. Fitton, 1975. A small label "praeter" (sic) J. F. P.

(Perkins). Female: Front wing 5.4 mm. Ovipositor 0.65 the length of front

wing. First gastral segment 4.5 x as long as wide apically.

Characteristics of the lectotype of Proclitus grandis: Labels: a Foerster

label Aachen, F, 23 gl. Lectotype label Townes, 1964. Female: Front wing

5.2 mm. Ovipositor 0.6 the length of front wing.

Characteristics of a paralectotype of Proclitus grandis: Labels: a

Foerster label Lousberg, 23.10. Var. 24 gl. Paralectotype label of the present

author. Male: Front wing 5.0 mm. Length of postannellus : length of second

flagellar segment = 4 : 3. Postannellus distinctly longer. First gastral segment

about 4.3 x as long as apical width.

Description of the female: Palpi, mandibles and clypeus yellow. Upper

tooth of mandible slightly longer. Clypeus rather flat, 2 x as wide as long,

with widely placed erect setae. Anterior tentorial pits impressed. Malar space

with a groove between eye margin and lateral corner of clypeus. Clypeus, face

and frons polished. Inner eye margins weakly converging towards clypeus.

Antenna yellow, the first segments long and slender. Pronotum polished, epomia

absent. Mesoscutum convex, polished, with sparse long hairs, notauli absent,

but proximally on margin indicated by a short carina and a dent. Scutellum

polished, with lateral carina at proximal corners. Propodeum polished. Basal

transverse carina absent, apical transverse carina well developed. Median longi-

tudinal and pleural carina present. Nervellus vertical not intercepted. Meso-

pleuron polished, prepectal carina not reaching front margin. Mesopleural pit

lying rather isolated. Sternaulus strong, but short. Legs, including coxae yellow,

long and slender. Apical halves of middle and hind femora somewhat expande d.

First tergite without glymma, about 4.3 x as long as wide apically. Median

dorsal carinae obsolete. All tergites polished, yellowish to brown. The second

tergite with thyridia. Ovipositor long, 0.5 - 0.7 of length front wing.

The male closely resembles the female. Length of postannellus : length

of second flagellar segment = 4: 3. First gastral segment 4.4 - 5.0 x as long

as apical width.

156 Contrib. Amer. Ent. Inst., vol. :20, 1983

Material examined: Austria: F M, Esternberg, 22-25.VII.1964 (coll. Zwart);

F, Poggstall (Nord O.) VII.1967 (coll. Zwart). Germany: F, lectotype

Proclitus grandis, Aachen; M, paralectotype P. grandis, Lousberg, 23.X (coll.

Foerster, Muenchen); F, Herrsching (Bayern), Widdersberg (Sumfiges Tal) 22.VI.

1974 (coll. Haeselbarth); M, Andechs (Ob. Bayern) 18.V.1959 (coll. Haeselbarth);

M, Zorneding (Ob. Bayern) 1.VI.1958 (coll. Haeselbarth); F, Wiesen (Spessart)

27.1X.1960; F M, Wiesen, 13.1X.1961 (coll. Haeselbarth); F, Lochmuhle (trap)

Nd Spessart, 17-22.1X.1971 (coll. van Rosseum); F, Lippoldshausen, 5.VI.1966 (coll.

Haeselbarth); F, Witzenhausen, 14.VIII.1965 (coll. Haeselbarth). Ireland: F, lecto-

type Proclitus praetor, no labels (coll. Haliday, Nat. Museum, Dublin).

Italia: M F, St. Peter (Ahrntal) (Sudtirol) 1300 m, 31.VII.1966 (coll. Haeselbarth);

F, Bolzano, 1914, leg. Smits van Burgst (coll. Agric. Univ. Wageningen); M,

Bolzano, Sarntal, 1250 m, 20.VI.1976 (coll. Zwakhals); F, Campi, Riva s. Garda,

1500 m, 7.1X.1967 (coll. Haeselbarth). Netherlands: M, Beetsterzwaag, 12.VI.

1967 (coll. Zwart); M, Ommen (Ov.) 5.VI.1964 (coll. Zwart); 1M, 4 M, Ede

(Geldl.) IX, X.1970 (coll. van Rossem); 2 F, Ede, 2.VII.1971 (coll. Zwakhals);

F, Vogelenzang (Nd Holl.) 30.1X.1933, leg. Barendrecht (coll. Zwart); 30 F M,

Asperen (Zd Holl.) VI,VII,VII,IX,X. 1972-1973 (coll. Zwakhals); F, Venlo. 19,VII.

1975, leg. van Aartsen (coll. Zwakhals); M, Maasbracht (L.) 13.V.1966, leg.

Lefeber (coll. Zwart). Sverige: 2 M, Orebro Lan, Klysna (Norrberga) 8.VII-

2.VI11.1979 (coll. van Rossem).

2. Proclitus comes (Haliday in Curtis)

Cryptus (Clepticus) comes Haliday in Curtis, 1838, Ann. Nat. Hist. 2:

116, F.

Proclitus macrurus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

283.116.

Proclitus macrurus: Aubert, 1977. Spixiana 1(2): 142.

Proclitus perditorius Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 116. (sec. Aubert, 1977). |

Proclitus caudiger Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 117. (sec. Aubert, 1977).

Proclitus pallens Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 117. (sec. Aubert, 1977).

Characteristics of the lectotype of Cryptus (Clepticus) comes: Labels:

lectotype label M. G. Fitton, 1975. A small label "comes" J. F. P. (Perkins).

Female: Front wing 4.5 mm. Ovipositor 0.77 of length front wing. First gastral

segment 2.4 x as long as wide apically.

Characteristics of the lectotype of Proclitus macrurus: Labels: a Foerster

label Aachen, F, 21 gl., a box label macrurus Frst. Lectotype label of the

present author. Female : Front wing 3.75 mm. Clypeus weakly convex, 2.3

x as wide as long. Apical margin of clypeus with a fringe of setae. Anterior

tentorial pits somewhat impressed, with a groove between eye margin and lateral

corner of clypeus. Ovipositor exceptionally long, 0.8 of length front wing.

Characteristics of the lectotype of Proclitus perditorius : No locality

label. Female: Ovipositor 0.76 of length front wing. Characteristics of a para-

lectotype: a Foerster label Aachen, F, head missing. Ovipositor 0.92 of length

front wing.

M= Male. F = Female.

van Rossem: Proclitus (Ichneumonidae) , 157

Characteristics of the lectotype of Proclitus caudiger: Labels: a Foerster

label Lousberg, 25.6. Ovipositor 0.83 of length front wing.

Characterics of the lectotype of Proclitus pallens: Labels: a Foerster

label Aachen. Female: Ovipositor 0.87 of length front wing.

Male: Unknown.

Material examined: Austria: 2 F, Tirol, Aschbach, 1400 m, 18-25.VIII.1975

(coll. Zwakhals); 2 F, Esternberg, 25-31.VII.1964 (coll. Zwart); F, Peilstein, 11.

VIII.1964, leg. den Hoed (coll. Zwart). Germany: F, lectotype of Proclitus

macrurus, Aachen (coll. Foerster, Muenchen); F, Glonn (Ob. Bayern), 18.VIII.

1970, Heidelbeere (coll. Haeselbarth); F, Harthausen bei Muenchen, 20.VIII.1969

(coll. Haeselbarth); F, Geisenfeld (Ob. Bayern), 16.VIII.1972, Heidelbeere (coll.

Haeselbarth); F, Andechs (Ob. Bayern), 10.X.1950 (coll. Haeselbarth). British

Isles: F, lectotype of Cryptus (Clepticus) comes , no labels (coll. Haliday,

Nat. Museum, Dublin). Italia: F, St. Peter (Ahrntal) (Sudtirol), 1600 m, 26.VII.

1967 (coll. Haeselbarth); F, Funes (Prov. Bolzano), 20.VII.-9.VIII.1968, Selva Near,

1400-2000 m (coll. van Rossem); F, Campi, Riva s. Garda, 1000 m. 7.VII.1966 |

(coll. Haeselbarth). Netherlands: F, Ede (Prov. Gelderl.) 20.1X.1970; F, Ede,

5.X.1970 (both coll. Zwakhals); F, Amsterdamse Waterleiding Duinen, 26.1X.1942,

leg. Barendrecht (coll. Zwart). Sverige: F, Dalarna, Transtrand, Hemfjall

Stangen, VII.1976 (coll. van Rossem).

The localities of the synonymous species are not repeated.

3. Proclitus fulvicornis Foerster

Proclitus fulvicornis Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 114.

Proclitus fulvicornis: Aubert, 1977. Spixiana 1(2): 144

Proclitus inquietus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 114. New synonym.

Proclitus cupidus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 114. New synonym.

Proclitus evacuator Foerster, 1871. Verh. Naturh. Ver. Preuss. Rbheinl.

28: 115. New synonym.

Proclitus evacuator: Aubert, 1977. Spixiana 1(2): 143.

Proclitus periculosus Foerster, 187]. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 115.

Plectiscus (Proclitus) heterocerus Thomson, 1888. Opusc. Ent. 38

(Fase: 12)¢' 1307.

? Proclitus ruficaudator Aubert, 1963. Bull. Soc. Ent. Mulh.:: 84.

Characteristics of the holotype of Proclitus fulvicornis: Labels: a

Foerster label Aachen, F, 17 gl. A box label fulvicornis Frst. Holotype label

of present author. Female: Front wing 3.5 mm. Clypeus polished, 2.4 x as

wide as long, with a conspicuous fringe of long setae. Malar space wide.

Anterior tentorial pits impressed, in the malar space limited by a strong carina

between eye margin and the lateral corner of the clypeus. Antenna short,

0.74 of length front wing. Median longitudinal carinae reduced to short stubs

on apical transverse carina. Legs, including coxae, yellow. Hind femur some-

what expanded, apical half 1.4 x as wide as base.

155 Contrib. Amer. Ent. Inst., vol. 20, 1983

First gastral segment 2.6 x as long as wide apically. Median dorsal carinae

present. Thyridia present, but weak. Second tergite fuscous, with yellow apical

margin. Third tergite yellow. Ovipositor 0.47 of length front wing.

Characteristics of the lectotype of Proclitus inquietus: Labels: a Foerster

label Lousberg, 26.10, 17 gl. Lectotype Aubert, 1967. Ovipositor 0.38 of length

front wing. The specimen is identical with P. fulvicornis.

Characteristics of the holotype of Proclitus cupidus: Labels: a Foerster

label Lousberg, 14.9, 17 gl. Holotype, no label. Ovipositor 0.37 of length front

wing. The specimen is identical with P. fulvicornis.

Characteristics of the holotype of Proclitus evacuator: Labels: a Foerster

label Lousberg, 25.10, F, 18 gl. A box label evacuator Frst. Holotype label

of the present author. Female: Front wing 4.75 mm. Clypeus convex, polished,

1.8 x as wide as long, apical margin straight, protruding. Anterior tentorial

pits impressed, in the malar space limited by a strong carina between eye mar-

gin and the lateral corner of the clypeus. Antenna short, 0.78 of length front

wing. Median longitudinal carinae strongly diverging. Hind femur weakly ex-

panded, apical half 1.5 x as wide as base. First gastral segment 2.5 x as long

as wide apically. Median dorsal carinae present. Thyridia rather strong and

long, limiting the proximal corners of the second tergite. Second tergite fus-

cous, with yellow apical margin. Third tergite yellow. Ovipositor 0.38 of

length front wing.

Characteristics of the lectotype of Proclitus periculosus: Labels: a

Foerster label Lousberg, 9.7. Lectotype label Aubert, 1967. Ovipositor of the

right specimen 0.46 of length front wing.

Characteristics of the holotype of Plectiscus (Proclitus) heterocerus

Thomson: Labels: P&l (P&lsjo, Skane), a Thomson label, "heterocerus", a

Thomson label. Holotype label of Townes, 1964. An Aubert label P-

fulvicornis Foerst.? The specimen is identified with P. fulvicornis Foerst.

but it has a much longer ovipositor, 0.7 of length front wing.

Characteristics of the female of Proclitus fulvicornis: Front wing

3.5 ~ 4.75 mm. Clypeus convex, protruding, the apical margin straight. An-

terior tentorial pits strongly impressed, with a carina between eye margin and

lateral corner of clypeus. The origin of the radius is from the distal 0.52 -

0.56 of the stigma. First gastral segment about 2.6 x as long as wide apically.

Median dorsal carinae present. Thridia present. Gastral tergites polished.

Ovipositor 0.36 - 0.70 of length front wing.

Male: Unknown.

Material examined: The localities of the type material of the synonymous

Foerster species are not recapitulated. Germany: F, holotype of P. fulvi-

cornis, Aachen, 17 gl. (coll. Foerster, Muenchen); F, Hochstadt (Bayern), 26.

VIII.1973 (coll. Haeselbarth); F, Hedemunden, 12.V.1966 (coll. Haeselbarth).

Italia: F, Sudtirol, Unser Frau in Schnals, 1600 m, 2.1X.1967 (coll. Haeselbarth);

F, Villnosz (Funes), 1400 m, 26.V1.1976 (coll. Zwakhals). Netherlands: 2 F,

Ede (Prov. Gelderl.), 15.X1.1970 (coll. Zwakhals); F, Asperen (Prov. Zd Holl.),

25.V1I.1972 (coll. Zwakhals). Sweden: F, holotype of Plectiscus (Proclitus)

heterocerus Thomson, P&l (P&lsjo, Sk&ne) (coll. Thomson, Lund.

van Rossem: Proclitus (Ichneumonidae) 159

4. Proclitus paganus (Haliday in Curtis)

Cryptus (Clepticus) paganus Haliday in Curtis, 1838. Ann. Nat. Hist.

2: 65 Fe.

Proclitus providus Foerster, 1871, Verh. Naturh. Ver. Preuss. Rheinl.

28: 114, F.

Proclitus providus; Aubert, 1977. Spixiana 1(2): 145.

Proclitus autumnalis Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 115. (sec. Aubert, 1977), F.

Proclitus curiosus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

20: 115. (sec, Aubert, 1977), F.

Proclitus instigator Foerster, 187]. Verh. Naturh. Ver. Preuss. Rheinl.

28: 115. (sec. Aubert, 1977), F.

Proclitus conturbator Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 115. (sec. Aubert, 1977), F.

Proclitus clypearis Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 115 and 117. (sec. Aubert, 1977), M, F.

Proclitus dimidiatus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 115. (sec. Aubert, 1977), F.

Plectiscus (Proclitus) longitarsis Thomson, 1888.

(Fasc. 12): 1306. (sec. Aubert, 1977), F.

Characteristics of the lectotype of Cryptus (Clepticus) paganus:

lectotype label M. G. Fitton, 1975.

Opusc. Ent. 38

Labels:

A small label "paganus" J. F. P. (Perkins).

Female: Front wing 4.0 mm. Ovipositor 0.30 of length front wing. Clypeus

flat, the apical margin arcuate. Anterior tentorial pits impressed and with

a carina between eye and clypeus.

Characteristics of the holotype of Proclitus providus: Labels: a Foerster

label Lousberg, 25.10, 18 gl., F. A box label providus Frst. Holotype label

of the present author. Female: Front wing 3.75 mm. Clypeus polished, flat,

2.1 x as wide as long, with an apical fringe of setae, the apical margin arcuate.

Anterior tentorial pits strongly impressed and in the malar space with a carina

between eye margin and lateral corner of clypeus. First gastral segment 2.3

x as long as wide apically. Median dorsal carinae present. Thyridia present.

Ovipositor short, 0.27 of length front wing.

Characteristics of the male paralectotype of Proclitus clypearis: The

male of P. providus was described by Foerster with the name P. clypearis.

There are 2 paralectotypes, both pinned on the same block. The right speci-

men was claimed to be P. providus by Aubert and I labelled this specimen

as the paralectotype. I nevertheless think that also the left specimen is P.

providus. Labels: a Foerster label Aachen, M, 20 gl.; an Aubert label. Para-

lectotype label of the present author (right specimen). Male: Front wing 4.1

mm. The anterior tentorial pits impressed and with a carina between the eye

margin and clypeus. The clypeus is flat, 2.5 x as wide as long.

Labels: a Foerster

Identical with P. providus.

Characteristics of the holotype of Proclitus autumnalis:

label Lousberg, 25.10, 18 gl., F. Ovipositor short.

Characteristics of the lectotype of Proclitus curiosus: Labels: a Foerster

label Lousberg, 25.10, F. Ovipositor short. Identical with P. providus . A

paralectotype, Lousberg, 25.10, 19 gl., F, identical with P. providus.

160 Contrib. Amer. Ent. Inst., vol. 20, 1983

Characteristics of the lectotype of Proclitus instigator: Labels: a

Foerster label Lousberg, 25.10, F. Ovipositor short. Identical with P.

providus. A paralectotype, Lousberg, 15.10, F, 19 gl., identical with

P. providus.

Characteristics of the lectotype of Proclitusconturbator: Labels: a

Foerster label Lousberg, 31.10, F. Identical with P. providus. A paralectotype,

Aachen, 19 gl., F. Ovipositor slightly longer, 0.30 of length front wing. A

second paralectotype, Lousberg, 25.10, F. Identical with P. providus.

Characteristics of the lectotype of Proclitus clypearis: Labels: a Foerster

label Lousberg, 31.10, F. Identical with P. providus. There are 6 paralectotype

females: Aachen, ovipositor 0.28 of length front wing; Lousberg, 31.10, oviposi-

tor 0.27; no label, ovipositor 0.24; Lousberg, 23.10, ovipositor 0.36; Lousberg,

31.10, ovipositor 0.22; Lousberg, 8.10, ovipositor 0.31.

Characteristics of the lectotype of Proclitus dimidiatus: Labels: a

Foerster label Lousberg, 15.10, ovipositor 0.28, identical with

A paralectotype, Lousberg, 27.10, 20 gl., F, ovipositor 0.31, identical with

P. providus.

Characteristics of the holotype of Plectiscus (Proclitus) longitarsis

Thomson. Labels: Rostock, 5.10.87, a Thomson label bongitarsus. Label of

Aubert Proclitus providus Foerster. Holotype label Fitton, 1978. Ovipositor

0.32 of length front wing. Anterior tentorial pits strongly impressed. Specimen

identical with P. providus.

Material examined: The localities of origin of the type material of the

synonymous species are not recapitulated. Ireland: F, lectotype of Cryptus

(Clepticus) paganus, Galway, Eyrecourt (coll. Haliday, Nat. Museum, Dublin).

Germany: F, holotype of P. providus, Lousberg (near Aachen), 25.10 (coll.

Foerster, Muenchen); 2 M, paralectotypes of P. clypearis, Aachen (coll.

Foerster, Muenchen); 3 F, Nord Spessart, Lochmuhle, 17-22.1X.1971, trap (coll.

van Rossem); 1 F, Wiesen, Spessart, 13.1X.1961 (coll. Haeselbarth). Netherlands:

1 M, Ede, 1.X1.1970 (coll. C. J. Zwakhals); 21 F, Ede, 1,8,15,22.X1.1970 (coll.

Corj. Zwakhaisie ll Py Bde BVI Ve23iV.,300V43 VIS VES2ZZVELI9TI (coll,

C. J. Zwakhals); 14 F, Asperen (Prov. Zd Hol.), the months V,VI,X and XI of

1972 and 1973 (coll. Zwakhals). Sverige: 1 F, Dalarna, Transtrand,

Hemfjaellstangen, VII.1976 (coll. van Rossem). Additional speciemsn from the

Netherlands: F, Amsterdam, X.1941, leg. Klaassen (coll. Zwart); 10 F,

Amsterdamse Bos, 2.XI.1952, leg. Wiering (coll. Zwart); 4 F, Burgst (Prov. Nd

Br.), X.1912, X.1913, X1.1913, II.1915 (coll. Smits van Burgst, Wageningen).

5. Proclitus zonatus (Gravenhorst)

Plectiscus zonatus Gravenhorst, 1829, Ichneumonologia Europaea 2(2):

984, F, no. 342. New combination.

Proclitus absconditus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 114.

Proclitus absconditus: Aubert, 1977. Spixiana 1(2): 146.

Proclitus contemptibilis Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 115. (sec. Aubert, 1977).

Proclitus punctatus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 115. (sec. Aubert, 1977).

van Rossem: Proclitus (Ichneumonidae) 161

Proclitus inferior Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 116. (sec. Aubert, 1977). 3

Proclitus displicitus Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 116. (sec. Aubert, 1977). |

Proclitus sordidus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 116. (sec. Aubert, 1977).

Proclitus humilis Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 117. (sec. Aubert, 1977).

Proclitus marginatus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

28: 117. (sec. Aubert, 1977).

Proclitus inaestimabilis Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 118. (sec. Aubert, 1977).

Proclitus denticulatus Foerster, 187]. Verh. Naturh. Ver. Preuss. Rheinl.

28: 118. (sec. Aubert, 1977).

Characteristics of the lectotype of Plectiscuszonatus: Labels: f an ori-

ginal Gravenhorst specimen. MZW 244. Female: Front wing 3.7 mm. Mandi-

ble tipe not twisted. Clypeus convex, polished, yellowish brown, 1.6 x as wide

as long. Malar space wide, with a groove. Anterior tentorial pits open. Face

polished, brown. Frons, vertex and gena polished. Antennae missing, except

for slender right postannellus. Epomia strong. Mesoscutum polished, notauli

present. Scutellum with carina to apex. Propodeum coriaceous, apical transverse

carina strong. Median longitudinal carina and basal transverse carina absent.

Propodeum and upper part of metapleuron with conspicuous long, erect hairs.

Nervellus vertical, discoidella absent. Mesopleuron polished, prepectal carina

to lower 0.3 of pronotum, not reaching the margin. Legs, including coxae,

yellow. First tergite rather wide, coriaceous, position of spiracles difficult

to determine as gaster is glued to thorax. Following tergites polished (diffi-

cult to see). Ovipositor 0.26 as long as front wing.

Characteristics of the holotype of Proclitus absconditus: Labels: a

Foerster label Koln, 25.9, F, 18 gl. A box label absconditus Frst. An Aubert

label: Plectiscidea tenuicornis Foerst. (= Proclitus absconditus Foerst.).

Female: Front wing 3.1 mm. Clypeus 1.3 x as wide as long. Malar space ex-

ceptionally wide, about 0.45 of width face, with a weak furrow. Anterior

tentorial pits not impressed. Gena wide, about 0.5 of width face. Epomia

present. Lateral carinae of scutellum running to apex, not meeting. Apex

of scutellum with longitudinal striation. Median longitudinal carinae diverging

proximally. Lateral longitudinal carina present. Hind femur not swollen. First

gastral segment 1.8 x as long as wide, coriaceous, median dorsal carinae absent.

Second tergite for the greater part coriaceous, apical margin polished, thyridia

absent. Ovipositor 0.26 of length front wing.

Characteristics of the male: The main characteristics of the male agree

with those of the female. Malar space and gena wide. Lateral carinae of

scutellum running to apex. Hind femur not swollen. First tergite coriaceous,

median dorsal carinae present.

Material examined: The localities of the type material of the synonymous

species of Foerster are not recapitulated. They all originated from Aachen

and Lousberg near Aachen (Germany).

No locality, F, holotype of Plectiscus zonatus (coll. Museum Zcologiczne,

Wroclaw). Germany: F, holotype of P. absconditus, Kony 2u1X4 Boek

162 Contrib. Amer. Ent. Inst., vol. 20, 1983

(coll. Foerster, Muenchen). Netherlands: F, Ede (Prov. Gelderl.), 28.1X.1970;

F, Ede, 17.X.1970; 5 F, Ede, 1.XI., 22.X1.1970; M, Noordwijk (Prov. Zd Hol.),

N. Leeuwenhorst, 22.X.1966, leg. van Ooststroom; F, Asperen (Prov. Zd Hol.),

25.X.1972; 3 M, Asperen, 24.VIII., 10.1X.,29.1X.1973 (all Dutch coll. Zwakhals).

6. Proclitus fulvipectus Foerster

Proclitus fulvipectus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

260 TD,

Proclitus fulvocingulatus Strobl, 1903. Mitt. Nat. Ver. f. Steierm.

40: 134. (sec. Aubert).

Proclitus fulvipectus: Aubert, 1977, Spixiana 1(2): 145.

Characteristics of the lectotype of Proclitus fulvipectus: Labels: no

locality label present, a box label fulvipectus Frst. The left specimen is the

lectotype, labelled by Aubert, 1977. Female: Front wing 3.6 mm. Clypeus

medially somewhat convex, about as wide as width face. Anterior tentorial

pits conspicuous, not impressed. There is a carina between eye margin and

lateral corner of the clypeus. Laterally of the carina a groove is present.

Right front wing missing. Propodeum with stubs of median longitudinal carinae

scarcely indicated. Hind femur weakly expanded, apical half 1.2 x as wide

as base. Right middle tarsus and right hind leg missing. First gastral seg -

ment 1.9 x as long as apically wide, median dorsal carinae present. Ovipositor

0.36 of length front wing.

Male: I did not see the male mentioned by Aubert (1977).

Material examined: Austria: F, lectotype of Proclitus fulvocingulatus,

Styriae alp. Strobl (? Wirtsgraben bei Hohentauern, August) (Strobl collection,

Admont, Austria). Germany: F, lectotype of Proclitus fulvipectus, no

locality label (coll. Foerster, Muenchen); F, Hann. Muenchen, A, 25.VI.1967

(coll. Haeselbarth, Muenchen).

7. Proclitus attentus Foerster

Proclitus attentus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

Zor il).

Proclitus attentus: Aubert, 1977. Spixiana 1(2): 143.

Proclitus gracilentus Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 117. (sec. Aubert, 1977).

Characteristics of the lectotype of Proclitusattentus : Labels: a Foerster

label Lousberg, 15.10. Mesoxanthus Frst. = fcssulatus Frst. = attentus Frst.

det. J. F. Aubert. A box label attentus Frst. Lectotype label of the present

author. Female: Front wing 4.0 mm. Clypeus convex, 2 x as wide as long,

polished. Malar space wide, 0.33 of width face, with a furrow. Face and frons

polished. Antenna short, 0.78 of length front wing. Hind femur expanded in

apical half, 2.2 x as wide as base. First gastral segment 3 x as long as wide

apically. Median dorsal carinae present. All tergites polished. Second tergite

with apical margin yellow, third tergite yellow. Thyridia weakly visible. Ovi-

positor 0.41 as long as front wing.

Characteristics of the lectotype of Proclitus gracilentus: Labels: a

Foerster label Aachen, M, 19 gl. A box label gracilentus Frst. I consider the

left specimen to be the lectotype. Male: Front wing 3.3 mm. Width of malar

space the same as width of mandibular base. Radius originating at distal

van Rossem: Proclitus (Ichneumonidae) 163

0.59 of stigma. The first gastral segment 3.0 x as long as width apically.

Hind femur expanded, apical half 1.7 x as wide as base.

Characteristics of the female of P. attentus : Front wing 3.2 - 4.5

Clypeus convex, protruding, the apical margin with a rather conspicuous fringe

of setae. Malar space wide, 0.33 of width face, with a furrow. Anterior ten-

torial pits not impressed. Radius originating at distal 0.54 of stigma. First

gastral segment about 3 x as long as wide apically. Ovipositor between 0.37 -

0.48 of length front wing.

Characteristics of the male of P. attentus: Front wing 3.2 - 4.0 mm.

Length of postannellus : length of second flagellar segment = Il : 10 (post-

annellus slightly longer). First gastral segment 3.4 - 3.8 x as long as

apically wide.

Material examined: Austria: M, Haus Gfohlalm, Heidelbeere, 1350 m,

28. VI. 1972 (coll. Haeselbarth). Helvetia (?): GR. M, Tarasp Avrona, 1450

m, 2. VIII. 1973, Erica, Bergfohrenw. (coll. Haeselbarth). Germany: F, lecto-

type of P. attentus, Lousberg (bei Aachen), 15.10 (coll. Foerster, Muenchen);

M, lectotype of P. gracilentus , Aachen (coll. Foerster, Muenchen); 2 F, Nord

Spessart, Lochmuhle, 17 - 22. IX. 1971, trap (coll. van Rossem); F M, Wiesen,

Spessart, 25. IX. 1960 (coll. Haeselbarth); F, Herrsching, Widdersberg (Bayern),

sumpfiges Tal, 22. VI. 1974 (coll. Haeselbarth); F, Rottenbuch (Bayern), 3. VI.

1974 (coll. Haeselbarth); M, Taufkirchen bei Muenchen, 26. VI. 1968 (coll.

Haeselbarth); F, Leppoldshausen, 21. VIII. 1966 (coll. Haeselbarth); 2 M,

Geierlambach, Heidelbeere, 7. VII. 1970 (coll. Haeselbarth); M, Bramwald, Nord

Sachsen, 30. VII. 1967 (coll. Haeselbarth). Netherlands: 2 F, Ede (Prov.

Gelderl.) 17. X. 1970 and 6. VII. 1971; F, Nunspeet (Prov. Gelderl.) 14. VI. 1975;

2 M, Overveen (Prov. Nd Hol.) 3l. V. and 19. X. 1974; 8 F, Asperen (Prov. Zd

Hol.) 28. 1%, 5. Mi 95.) Xo 199225. W110, Xi 107303 MW, Aeperem. (Prove Zd

Hol.) 1. VIII. 1972; 2. VIII. 6. VIII. 1973 (all Dutch specimens from coll.

Awakhals). Additional specimens from the Netherlands: 4 F, 4 M, Bakkum-

merduin (Prov. Nd Hol.), ex Armillaria mellea (Vahl ex Fr.) Kumm. 27. IX.

1931, leg. Sorgdrager and Westenberg (coll. Zwart); F, Liesbosch (Prov. Nd Br.),

V. 1913 (coll. Smits van Burgst, Wageningen).

8. Proclitus rudis Foerster

Proclitus rudis Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl. 28:

115.

Proclitus rudis: Aubert, 1977. Spixiana 1(2): 144.

Characteristics of the holotype of Proclitus rudis: Labels: a Foerster

label Aachen, F, 19 gl. A box label rudis Frst. Holotype label of the present

author. Female: Front wing 4.3 mm. Clypeus polished, about 3 x as wide

as long and with rather conspicuous long setae. Malar space with a groove.

Width of malar space about equal to width of mandibular base. Anterior

tentorial pits not impressed. Face and frons polished. Occipital carina some-

what lamelliform. Postannellus and the following segment of the same length.

Antennae broken off. Occiput steep, somewhat concave directly behind ocelli.

Weak epomia present. median longitudinal carinae long, reaching proximal

margin of propodeum, somewhat diverging proximally. Radius originating at

about 0.50 of stigma. Hind femur somewhat expanded, apical half 2 x as wide

as base. First gastral segment in comparison with other species short and

164 Contrib. Amer. Ent. Inst., vol. 20, 1983

wide, 2 x as long as wide apically. First tergite with longitudinal striation,

median dorsal carinae present. The gaster beyond segment two rather com-

pressed. Ovipositor 0.39 of length front wing.

Material examined: Germany: F, holotype of Proclitus rudis, Aachen

(coll. Foerster, Muenchen). Austria: 2 F, Karnten, Bodental, 1100 m, 29. VI.

and 14. VII. 1981 (coll. Zwakhals, Arkel).

9. Proclitus subsulcatus Foerster

Proclitus subsulcatus Foerster, 1871. Verh. Naturh. Ver. Preuss.

Rheinl. 28: 117.

Characteristics of the holotype of Proclitus subsulcatus: Labels: a

Foerster label Aachen, M, 20 gl. A box label subsulcatus Frst. Holotype label

of the present author. Male: Front wing 4.87 mm. Face wide, about 0.45

of frontal width of head including eyes, with a row of conspicuous long, erect

setae along inner margin of eye. Clypeus exceptionally wide, almost 3 x as

wide as long, the apical margin arcuate. Lateral corner of clypeus reaching

beyond line of inner margin of eye. Clypeus with widely placed long setae.

Anterior tentorial pits open, not impressed. Malar space narrow, about 0.5

of width mandibular base. Scape somewhat enlarged, oval in shape. Median

longitudinal carinae poorly developed, strongly diverging. Legs, including coxae

yellow. Middle and hind coxae and trochanters, including hind femur, with

conspicuous, subadpressed setae. Hind femur somewhat swollen, apical half

about 1.6 x as wide as base. First gastral segment 2.8 x as long as apically

wide. Median dorsal carinae not running beyond spiracles. Thyridia rather

large, but connection between first and second segment somewhat damaged.

All tergites with conspicuous long subadpressed setae.

I did not succeed in finding a matching female.

Material examined: Germany: M, holotype, Aachen (coll. Foerster,

Muenchen).

10. Proclitus albidipes Foerster

Proclitus albidipes Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

Zor 7M,

Proclitus bicarinatus Foerster, 1871. Verh. Naturh. Ver. Preuss. Rheinl.

2B Th os

Characteristics of the holotype of Proclitusalbidipes: Labels: a

Foerster label Lousberg, 18.9, 18 gl. A box label albidipes Frst. Holotype

label Aubert, 1967. Proclitus periculosus Foerst. = albidipes Foerst. det.

J. F. Aubert. Male: Front wing 3.5 mm. Clypeus convex, polished, 2.2 x as

wide as long, with long setae. Malar space not developed, eye margin almost

touching mandibular base. Anterior tentorial pits not impressed, open. Face

polished. Inner eye margins weakly converging. Width of face at antennal

sockets 1.4 x as wide as width at clypeal margin. Length of postannellus :

length of second flagellar segment = 13 : 10. Mesoscutum polished, notauli

reduced to a dent at the margin and showing a short carina which is turned

inward. Propodeum with apical transverse carina strong. Lateral longitudinal

carinae obsolete. Mesopleuron with prepectal carina running up to about 0.5

of pronotal margin, not reaching the margin. First gastral tergite with

van Rossem: Proclitus (Ichneumonidae) 165

median dorsal carinae present and strong dorsolateral carina beyond the

spiracle. Second tergite with some longitudinal sculpture. The following ter-

gites polished.

Characteristics of the holotype of Proclitus bicarinatus: Labels: a

Foerster label Aachen, M, 19 gl. A box label bicarinatus Frst. Holotype

label of the present author. Male: Front wing 3.7 mm. Length of postan-

nellus : length of second flagellar segment = 12 : 10 (6: 5). Inner eye margins

weakly converging, width of face at antennal sockets 1.4 x as wide as width

at clypeal margin. Notauli reduced to a dent at the margin and with the

marginal carina turned inward. First gastral tergite with median dorsal carinae

present and strong dorsolateral carina beyond the spiracle.

Female: Unknown. Aubert (1977) considered this male to belong to the

female of P. fulvicornis. I do not see a motive to follow this, especially

not as the anterior tentorial pits of the P. fulvicornis female are strongly

impressed and with a conspicuous carina between eye margin and clypeus.

The malar space of P. albidipes is not developed.

Material examined: Germany: M, holotype of Proclitus albidipes ,

Lousberg (near Aachen), 18. IX (coll. Foerster, Muenchen); M, holotype of

Proclitus bicarinatus, Aachen (coll. Foerster, Muenchen); M, Glonn (Ober

Bayern), 18. VIII. 1970, Heidelbeere; M, Hann. Muenchen, 13. VIII. 1965 (both

specimens coll. Haeselbarth, Muenchen).

ACKNOWLEDGMENTS

The Director of the Zoologische Staatssammlung at Muenchen, Dr. E. J.

Fittkau, kindly permitted the author to study the Foerster Types in that

Institute. I also wish to thank Mr. E. Diller for his kind assistance and

hospitality. The following persons loaned interesting material from their col-

lections: Dr. E. Haeselbarth, Muenchen; Dr,. C. J. Zwakhals, Arkel (Nether-

lands). Prof. Dr. habil. G. Morge, Eberswalde-Finow (DDR), the curator of

the Strobl collection at Admont (Austria) and Drs. K. W. R. Zwart, the curator

of the Smits van Burgst collection which is in the custody of the Agricultural

University at Wageningen, allowed me to study respectively the Strobl and Smits

van Burgst collection. Dr. J. P. O'Connor kindly sent the Haliday Types which

are kept in the National Museum of Ireland. Dr. R. Danielsson of the Zoologi-

cal Museum at Lund gave me the possibility to examine the Thomson speci-

mens.

The author is indebted to the Uyttenboogaart-Eliasen Stichting at

Amsterdam for subsidy to travel to Muenchen.

TWO NEW PALEARCTIC SPECIES OFSESIOPLEX VIERECK

(HYMENOPTERA : ICHNEUMONIDAE) INCLUDING PHYLOGENY,

ZOOGEOGRAPHY AND A KEY TO THE WORLD SPECIES

P. M. Sanborne

Department of Biology, McMaster University

Hamilton, Ontario, L8S 4K1

Abstract

Two new Palearctic species of Sesioplex Viereck (Hym.: Ichneumonidae),

brunipalpus, n. sp., and cribus, n. sp., are described and illustrated. Two

Palearctic species, cerophagus (Grav.), and punctulatus Horstmann, are re-

moved from Sesioplex and placed in Campoplex [C. cerophagus Grav., C.

punctulatus (Horst.), n. comb.], Reasons for this are given. The generic

limits of Sesioplex are redefined, a key to the world species is provided, and

its phylogeny and zoogeography are discussed.

INTRODUCTION

I recently revised the North American species of Sesioplex Viereck

(Hymenoptera: Ichneumonidae) (Sanborne 1981). The primary diagnostic

character of the genus is a slender ovipositor (its depth at midlength distinct-

ly less than width of hind basitarsus) which bears a pronounced nodus im-

mediately in front of the dorsal notch. I have since discovered 2 Palearctic

species which possess identical ovipositors and agree in other characters as

well (see discussion following phylogeny and generic description). These

species, brunipalpus, n. sp., and cribus, n. sp. are placed in Sesioplex , de-

scribed and illustrated. The genus is redefined, its phylogeny and zoogeo-

graphy are discussed, and a key to the world species is provided.

Horstmann (1977) placed 2 European species, cerophagus (Grav.) and

punctulatus Horst., in Sesioplex and these were re-examined. They clearly

do not agree with Sesioplex (sensu Viereck). In my opinion, both belong

in Campoplex (sensu lato) and therefore, cerophagus (Grav.) is reinstated in

Campoplex (i.e., C. cerophagus Grav.) and punctulatus Horst. is placed in

Campoplex as a new combination [i.e., C. punctulatus (Horst.) n. comb.] (see

discussion following generic description).

MATERIALS AND METHODS

Only the holotypes of cribus, n. sp. and brunipalpus, n. sp. are avail-

able for study. They were sent to Dr. Klaus Horstmann (Zoologisches

Insitut, Wurzburg), an authority on Palearctic Porizontinae (= Campopleginae),

to establish whether or not the species had previously been described and

placed in other genera, and to try and locate additional material. He advised

me (personal communication, 1982) that both species were unfamiliar to him.

Their distinctiveness, and the need to provide additional evidence in support

of maintaining separate generic status for Sesioplex , prompted me to describe

the species at this time. Other material examined includes the lectotype of

Omorgus picticrus Thomson [= Sesioplexcerophagus (Gravenhorst)] and the

holotype of Sesioplex punctulatus Horstmann. The lectotype of Campoplex

cerophagus Grav. is housed in Breslau (Wroclaw), Poland and was unavailable.

166

Sanborne: Sesioplex (Ichneumonidae) 167

Terminology; ‘The terminology used in this paper is that of Townes

(1969) except that 'gena’ is used instead of 'cheek'.

Measurements: Some of these are described in my earlier paper

(Sanborne 1981). Additional measurements are made in the following manner.

The length of the areola is measured thorugh its middle from the apical mar-

gin of the basal area to a line defined by the junction of the median longi-

tudinal carinae with the basal transverse carina. The length of the petiolar

area is measured from this line, medially through the propodeal trough, to the

articulating membrane at the junction of the propodeum and abdomen. The

width of the areola is measured between the median longitudinal carinae at

their junction with the basal transverse carina. The width and height of the

petiole of the Ist abd. segment are measured at the midlength of the petiole.

The length of the petiole is measured, in lateral view, between the posterior

margin of the articulating membrane and the midpoint of the spiracle. The

post petiole length is measured, in the same view, from the midpoint of the

spiracle to the middle of the apical margin of the postpetiole. The width of

the postpetiole is measured at its widest point. The length of the hind tarsal

claw is measured, in dorsal view, from the base to the apex and does not ac-

count for curvature of the claw. The empodium comprises a sclerotized basal

portion and a membranous apical portion. The apical margin of the sclerotized

base is indented medially so that there are 2 short lateral projections. The

length of the empodium is measured from the basal margin to the apical margins

of the lateral projections. The length of the ovipositor is measured relative

to the lengths of the hind femur and the thorax. For convenience, the length

of the thorax is measured, in lateral view, from the anterior (closest to head)

margin of the tegula to the apex of the propodeum. The thorax appears to

remainunmodified in spite of variation between species in the length and depth

of the hind femur and ovipositor. This becomes useful in resolving situations

where, for example, 2 species, one with a short hind femur and short oviposi-

tor, the other with a long hind femur and long ovipositor, give approx. the

same length : length ratios. It is possible to get different ratios when their

Ovipositors are measured relative to the length of the thorax. I have found

examples of this in current studies of the related genus Sinophorus. The tip

of the dorsal valve of the ovipositor consists of the portion from, but not in-

cluding, the dorsal notch to the apex of the valve and is measured, in lateral

view, along the dorsal surface (exclusive of claw) from the apical margin to

the point where the segment begins to constrict near the point of articulation

with the 4th segment.

Illustrations: All drawings were made with the aid of a camera lucida

attached to a Wild M5 dissecting microscope.

PHYLOGENY

The genus Sesioplex is considered a monophyletic group based on the

structure of the ovipositor which is very slender and possesses a pronounced

nodus in front of the dorsal notch. In addition, the suture separating the ter-

gite from the sternite of the petiolar segment of the abdomen is distinctly

below the midheight of the petiole. The genus appears to be closely allied

to Sinophorus and also appears to have affinities to Campoplex (see discussion

following generic description).

168 Contrib. Amer. Ent. Inst., vol. 20, 1983

The North American species (depressus Viereck, californicus Sanborne)

appear to be more closely related to each other than to either brunipalpus,

n. sp. or cribus, n. sp. from the Palearctic region. Both North American

species have a more or less spherical head with a broad face, perpendicular

2nd recurrent vein in the fore wing, hind femur approx. 3.8 - 4.0 x as long

as deep, and a straight ovipositor which is approx. 2.0 x as long as the hind

femur. They differ strongly in the structure of the mandible, petiole of the

Ist abd. seg. and color as described in my 1981 paper. I consider them to be-

long to the same species-group but do not consider them as sister-species.

Conversely, brunipalpus, n. sp. and cribus, n. sp. appear to be more

closely related to each other than to either of the North American species.

Nevertheless, the species are very distinct from one another and are not con-

sidered to be sister-species and may not even belong to the same species-

group. This may become clearer when the extent of intraspecific variation

is known. The lower margins of the cmpd. eyes are weakly convergent in

brunipalpus, n. sp., but weakly divergent in cribus, n. sp. (Figs. 1, 9). There

are strong differences in the punctation, wrinkling and depth of the propodeal

trough (Figs. 3, 12). The petiole of cribus, n. sp. is distinctly wider than

deep (Figs. ll, 13) while that of brunipalpus, n. sp. is approx. as wide as deep

(Figs. 4, 5). Features common to both species include a more or less triangu-

lar head (Figs. 1, 9), sloping 2nd recurrent vein in the fore wing, hind femora

of similar shape and dimensions (approx. 4.1 - 4.3 x as long as deep) and

petiole with a short baso-lateral groove (more pronounced in cribus,n. sp.).

The first three character states may represent the plesiomorphic condition and,

as such, cannot be used to establish a sister relationship.

In my opinion, brunipalpus, n. sp. is the most generalized of the 4

species discussed. Based on out-group comparisons with the apparently related

Sinophorus and Campoplex, the following character states are postulated to

be plesiomorphic: eye margins moderately notched opposite antennal sockets;

lower margins of cmpd. eyes weakly convergent; head more or less triangular

in frontal view; temples, in dorsal view, moderately sloping inwardly; propodeal

trough moderately excavated, with transverse wrinkling on the petiolar area;

hind femur 4.1 - 4.3 x as long as deep; front wing with 2nd recurrent vein

moderately sloping (approx. 20 - 30 degrees from vertical); hind wing with

nervellus intercepted near basal 0.2 by discoidella; ovipositor curved upward

(to what degree, i.e., weakly mod., strongly, is plesiomorphic has not been de-

termined). The following color character states are postulated to be plesio-

morphic: head, thorax, and abdomen entirely black (exclusive of mandibles,

palpi, tegula and abdominal sternum); hind tibia with distinct sub-basal and

apical dark bands, and median band white externally.

ZOOGEOGRAPHY

In my 1981 paper I suggested that the discovery of additional species might

shed light on the probable center of origin and pattern of speciation within

Sesioplex. The latter cannot be determined because of the apparent lack of

sister-pairs within the group. With regards to center or origin, I follow the

school of thought which says that the more primitive extant members of a

group will usually be found nearer to the area occupied by its ancestors than

the more derived members (in Pielou 1979, Wiley 1981). If this is correct and

Sanborne: Sesioplex (Ichneumonidae) 169

brunipalpus, n. sp. is the most primitive extant member of Sesioplex , then,

the genus has its center of origin in the Palearctic region. A more precise

location awaits the determination of the center of concentration of bruni-

palpus, n. sp. or the discovery of an even more primitive species.

Genus SESIOPLEX Viereck

Campoplex subg. Sesioplex Viereck, 1912. Proc. U.S. Natl. Mus. 42: 644.

Type-species: Campoplex (Sesioplex) depressus Viereck. Orig. desig.

Head, in frontal view, broadly triangular to spherical; approx. 1.9 - 2.5

as wide as combined height of face and clypeus. Compound eyes with margins

weakly to moderately notched opposite antennal sockets, weakly divergent to

weakly convergent below. Temples, in dorsal view, moderately sloping inward-

ly; in lateral view, approx. 0.8 as long as cmpd. eyes. Gena approx. 1.1 - 1.4

as long as basal width of mandible (measured to lower margin of mandible).

Genal carina joining oral carina at or above base of mandible. Thorax with

mesopleural suture impressed as a sharp groove. Propodeum with areola and

petiolar area completely confluent and forming a mod. broad to broad, weakly

to moderately excavated trough. First, 2nd, 3rd lateral areas usually completely

enclosed by carinae (costula often obsolete around lateral border of Ist lateral

area). Areola approx. 0.7 ~ 0.9 as wide and 0.3 as long as petiolar area and

approx. 0.4 as wide as distance between propodeal spiracles. Front wing approx.

3.8 - 7.8 mm; areolet present; 2nd recurrent vein mod. sloping to perpendicu-

lar. Hind wing with nervellus not intercepted or intercepted near basal 0.2

by discoidella which is in the form of a faint furrow. Discoidella strongly

arching in front of midlength. Hind femur approx. 3.8 - 4.3 as long as deep.

Hind tarsal claw with 5 - 7 pectins which do or do not extend to base of tip.

First abd. tergite with or without a lateral longitudinal groove. Suture separ -

ating Ist abd. tergite from sternite approx. 0.4 - 0.7 below midheight of pe-

tiole. Petiole as wide as or wider than deep. Apex of male clasper rounded.

Ovipositor approx. 2.0 ~ 2.4 as long as hind femur, with a pronounced nodus

immediately in front of dorsal notch, weakly upcurved to Straight, its depth

at midlength distinctly less than width of hind basitarsus. Tip of dorsal valve

of ovipositor approx. 0.8 - 1.1 as long as Sth hind tarsal segment (exclusive

of claw).

Discussion: In my 1981 paper I stated that Sesioplex appears to be

Closely allied with Sinophorus with affinities to Campoplex. This view has not

changed but the discovery of 2 new Palearctic species of Sesioplex and the

results of current studies of Sinophorus make it necessary to modify and ex-

pand some of the statements made in that paper.

The North American species of Sesioplex (depressus Viereck, cali-

fornicus Sanborne) possess a perpendicular 2nd recurrent vein in the fore wing

which I initially treated as a diagnostic feature of the genus. The Palearctic

species of Sesioplex (brunipalpus, n. sp.,cribus, n. sp.) have the 2nd re-

current vein weakly to moderately sloping respectively, as in the majority of

Sinophorus. The majority of Campoplex have the 2nd recurrent vein strongly

sloping (30 - 35 deg. from vertical). The hind wings of Sesioplex and

Sinophorus have the discoidella moderately to strongly arching in front of the

midlength (toward wing base) but the majority of Campoplex have the

170 Contrib. Amer. Ent. Inst., vol. 20, 1983

discoidella straight or weakly arching and positioned closer to the lower mar-

gin of the wing. This is the most consistent character that I have found for

separating Sinophorus and Sesioplex from Campoplex. However, it is not 100%

efficient as there are a few species of Campoplex which have the discoidella

moderately to strongly arching. The areolet in the fore wings of Sinophorus

and Sesioplex is large (except one undescribed species of Sinophorus and a

few specimens of Sesioplex depressus). The areolet of Campoplex is common-

ly small but there is overlapping with Sinophorus and Sesioplex.

The propodeal trough of Sesioplex is very similar to that of Sinophorus

with regards to the degree of excavation and width of the areola. In

Sesioplex and approx. 95% of Sinophorus, the areola is approx. 0.4 x as wide

as the distance between the propodeal spiracles, while in the majority of

Campoplex it is only 0.3 x as wide.

The genus Sesioplex differs from all Sinophorus and Campoplex in the

possession of a pronounced nodus immediately in front of the dorsal notch of

the ovipositor. The length of the ovipositor is an artificial but, nevertheless,

useful character in separating Sinophorus and Sesioplex. from most Campoplex.

It rarely exceeds 2.5 x the length of the hind femur in Sinophorus, whereas,

it almost always does so in Campoplex (often greatly). In Sesioplex it varies

from approx. 2.0 - 2.4 x the length of the hind femur. The depth of the ovi-

positor is another useful character. In Sinophorus, its depth at midlength is

rarely less than the width of the hind basitarsus, almost always less than in

Campoplex and, with present knowledge, always less than in Sesioplex. The

length of the tip of the dorsal valve in Sinophorus ranges from 1.0 - 2.0 x

the length of the 5th seg. of the hind tarsus (exclusive of claw) (usually about

1.3). It ranges from 0.5 - 1.0 x in Campoplex, with most species around 0.8 x.

This character is intermediate in Sesioplex and ranges from 0.8 - 1.1 x the

length of the 5th seg.

The 2 species placed in Sesioplex by Horstmann (1977), cerophagus

(Grav.), punctulatus Horst., are removed from the genus for the following

reasons: both species lack a pronounced nodus in front of the dorsal notch

of the ovipositor; areola of both species approx. 0.3 x as wide as distance be-

tween propodeal spiracles (as in Campoplex ); discoidella weakly arched in

cerophagus, mod. arched in punctulatus (within range of Campoplex ); tip of

dorsal valve of ovip. as long as 5th seg. of hind tarsus in cerophagus, approx.

0.7 x as long in punctulatus (as in Campoplex); clypeus small, strongly convex

in cerophagus (common in Campoplex, not found in Sesioplex, very rare in

Sinophorus ). On the basis of these characteristics, cerophagus is reinstated

in Campoplex (i.e., C. cerophagus Grav.) and punctulatus is placed in

Campoplex as a new combination [i.e.,Cc. punctulatus (Horst) n. comb.]

ARTIFICIAL KEY TO THE WORLD SPECIES OF SESIOPLEX

1 Abdominal tergites | - 6 partially to entirely ferruginous. Mandible with

upper tooth longer than lower tooth. First abdominal segment with

a distinct lateral longitudinal groove along the entire length of petiole.

Eastern United States and Canada .......... depressus Viereck

Abdominal tergites entirely black. Mandible with upper tooth longer or

shorter than lower tooth. First abdominal tergite without a lateral

longitudinal groove or with short baso-lateral groove ........ 2

Sanborne: Sesioplex (Ichneumonidae) 171

y Mandible with upper tooth shorter than lower tooth. First abdominal ter-

gite without a lateral groove. Ovipositor approx. 2.0 as long as hind

femur. “Cantormia oe! 2 Po ee es californicus Sanborne

Mandible with upper tooth slightly to distinctly longer than lower tooth.

First abdominal tergite with short baso-lateral groove. Ovipositor

approx. 2.2 ~ 2.4 x as long as hind femur. Palearctic region... 3

4 Mandible with upper tooth distinctly longer than lower tooth. Ovipositor

approx. 2.2 x as long as hind femur. Propodeal trough moderately ex-

cavated, few distinct punctures evident and with distinct transverse

wrinkling:..on PpeLioOlat AfCd inet oie WS Spo ees eee brunipalpus, nn. sp.

Mandible with upper tooth slightly longer than lower tooth. Ovipositor

approx. 2.4 x as long as hind femur. Propodeal trough shallow, with

dense punctures over entire surface and irregular, longitudinal wrinkles

OR OC MIG ALOR i sil ce ee sw hs ce ee ee cribus, n. sp.

1. Sesioplex brunipalpus, n. sp. (Figs. 1-8)

Diagnosis: Head approx. 2.0 as wide as height of face and clypeus. Man-

dible with upper tooth distinctly longer than lower tooth. Mesopleuron with

moderate size punctures (approx. .002 mm) on lower 0.5 which are separated

by distance approx. equal to dia. of single puncture. Hind tarsal claw with

pectins which extend to base of tip. First abd. tergite with petiole as wide

as deep. Ovipositor approx. 2.2 as long as hind femur, weakly upcurved. Hind

wing with nervellus weakly intercepted near basal 0.2 by discoidella which is

in form of faint furrow. Hind tibia with distinct sub-basal, apical dark bands.

Holotype female: Head approx. 2.0 as wide as height of face and clypeus

combined, triangular in frontal view. Compound eyes with margins mod. notched

opposite antennal sockets, weakly convergent below. Antennal flagellum with

35 ~ 40 articles (estimated, tips broken). Face strongly shagreened, matte,

mod. rugose medially, with large punctures which are separated by distance

less than dia. of single puncture. Clypeus strongly shagreened, matte, with

numerous punctures; distinctly separated from face by broad shallow impres-

sion; mod. convex, with antero-median field flattened; anterior margin mod.

convex, broadly, weakly produced medially. Anterior 0.4 of clypeus produced

below lower margin of cmpd. eye. Mandible with upper tooth distinctly longer

than lower tooth. Gena approx. 1.1 as long as basal width of mandible. Tem-

ple, in lateral view, approx. 0.8 as long as cmpd. eye; in dorsal view, mod.

sloping inwardly. Mesopleuron with coarse wrinkling anterior to speculum which

does not extend to mesopleural fovea. Specullum with numerous punctures

dorsally. Mesopleural fovea deeply impressed, elongate-oval. Lower 0.5 of

mesopleuron mod. shagreened, weakly polished, with med. size punctures (ap-

prox. .002 mm) which are separated by distance approx. equal to dia. of single

puncture. Propodeum with basal transverse carina mod. strong, forming 3-sided

border around areola (median length approx. 0.5 as long as lateral length).

Costula as strong as basal transverse carina along border with 2nd lateral area

but absent along lateral border of Ist lateral area. Median longitudinal carina

weak, highly irregular near and not distinctly joining basal transverse carina;

mod. strong, more or less straight medially. Lateral longitudinal carina highly

irregular, indistinct. First lateral area sparsely punctate. Propodeal trough

mod. excavated. Areola moderately shagreened, polished, with several distinct

172 Contrib. Amer. Ent. Inst., vol. 20, 1983

punctures and traces of transverse wrinkling apically; approx. 0.9 as wide and

0.3 as long as petiolar area and 0.4 as wide as distance between propodeal

spiracles. Petiolar area with approx. 15 mod. coarse transverse wrinkles. Hind

femur approx. 4.3 as long as deep. Hind tarsal claw approx. 1.7 as long as

empodium (estimated from middle claw, hind claws lost) with 5 - 6 strong,

mod. inclined pectins which extend to base of tip. First abd. tergite a small

baso-lateral groove, the sides of the tergite otherwise weakly, irregularly ru-

gose. Suture separating Ist abd. tergite from sternite approx. 0.4 below. mid-

height of petiole. Petiole as wide as deep and approx. 1.1 as long as post-

petiole. Postpetiole approx. 3.6 as wide as petiole, with mod. dense dorsal

pubescence. Ovipositor approx. 2.2 as long as hind femur and 1.8 as long as_

thorax, weakly upcurved, its depth weakly increasing from base to tip, its

depth at midlength distinctly less than width of hind basitarsus. Dorsal valve

of ovipositor with a pronounced nodus immediately in front of dorsal notch.

Tip of dorsal valve approx. as long as 5th seg. of hind tarsus (exclusive of

claw). Front wing approx. 6.1 mm, with 2nd recurrent vein mod. sloping.

Hind wing with nervellus straight, erect, weakly intercepted near basal 0.2 by

discoidella which is in form of weak furrow.

Color: Head, base, apex of mandible, thorax, abdomen, coxae, basal seg.

of foretrochanter, entire mid, hind trochanters (except apical margins), sub-

basal, apical bands of hind tibia, apical 0.6 of hind basitarsus, black. Median

area of mandible yellowish. Maxillary palpus with apical 4 segments fulvo-

ferruginous, basal seg. brown. Labial palpus brown. Tegula yellowish-white.

Apex of basal seg. and entire apical seg. of foretrochanter, apical margins of

mid, hind trochanters, femora, internal surfaces of all tibiae (except extreme

bases which are whitish), sub-basal, apical patches (external) of midtibia, ferru-

ginous. External surface of foretibia, externo~median surfaces of mid, hind

tibiae, basal 0.4 of hind basitarsus whitish.

Male: Unknown.

Holotype: Female, USSR. Kurovsk region, near Ramensk, 24. VIII. 1962,

Kameisova. Deposited in the Academy of Sciences, Leningrad.

Etymology: The species is named for the brown labial palpus.

Host and Biology: The holotype was taken on sugar beet. The host may

be Loxostege sticticalis (Linn.) (Pyralidae), a common pest of this plant.

2. Sesioplex cribus, n. sp. (Figs. 9 - 16)

Diagnosis: Head approx. 1.9 as wide as height of face and clypeus com-

bined. Mandible with upper tooth slightly longer than lower tooth. Lower

0.5 of mesopleuron with large punctures (approx. .004 mm) which are separ-

ated by distance approx: equal to dia. of single puncture. Propodeal trough

broad, shallow, with entire surface densely punctate. Hind tarsal claw with

pectins distributed along basal 0.5. First abd. tergite with petiole approx. 1.5

as wide as deep. Ovipositor approx. 2.4 as long as hind femur. Hind wing

with nervellus not intercepted by discoidella. Hind tibia with sub-basal band

indistinct.

Holotype female: Head approx. 1.9 as wide as face and clypeus combined,

broadly triangular in frontal view. Compound eyes with margins weakly notched

opposite antennal sockets, weakly divergent below. Face strongly shagreened,

Sanborne: Sesioplex (Ichneumonidae) 173

matte, mod. rugose, with large punctures (approx. .004 mm) which are separ-

ated by distance less than dia. of single puncture. Clypeus strongly sha-

greened, matte, with numerous punctures; distinctly separated from face by

broad, shallow impression; moderately, evenly convex, with anterior margin

mod. convex and not produced medially. Anterior 0.4 of clypeus produced be-

low lower margin of compound eye. Mandible with upper tooth slightly longer

than lower tooth. Gena approx. 1.2 as long as basal width of mandible. Tem-

ple, in lateral veiw, approx. 0.8 as long as cmpd. eye; in dorsal view, mod-

erately sloping inwardly. Mesopleuron with mod. coarse wrinkling anterior to

speculum which nearly reaches mesopleural fovea. Speculum with numerous

punctures dorsally. Mesopleural fovea deeply impressed, elongate, irregular.

Lower 0.5 of mesopleuron with large punctures (approx. .004 mm) which are

separated by distance less than dia. of single puncture. Propodeum with basal

transverse carina moderately strong, forming rounded, V-shaped border around

areola. Costula as strong as basal transverse carina along border of 2nd lateral

area, absent along lateral border of Ist lateral area. Median longitudinal

carina indistinct near junction with basal transverse carina, mod. strong, weak-

ly arching medially. Lateral longitudinal carina weak; highly irregular. First

lateral area densely punctate. Propodeal trough broad, shallow, with entire

surface densely punctate, the punctures large. Areola approx. 0.8 as wide,

0.3 as long as petiolar area and 0.4 as wide as the distance between the pro-

podeal spiracles. Hind femur approx. 4.1 as long as deep. Hind tarsal claw

approx. 1.5 as long as empodium, with 6 mod. strong, mod. inclined pectins

which extend along basal 0.5 of claw. First abd. tergite with a short, baso-

lateral groove. Suture separating Ist abd. tergite from sternite approx 0.5 be-

low midheight of petiole. Petiole approx. 1.5 as wide as deep, 1.4 as long as

postpetiole. Postpetiole approx. 3.1 as wide as petiole, with mod. dense dorsal

pubescence. Ovipositor approx. 2.4 as long as hind femur, 1.9 as long as thorax,

straight, weakly increasing in depth from base to tip; depth at midlength

slightly less than width of hind basitarsus. Dorsal valve with a pronounced

nodus in front of dorsal notch. Tip of dorsal valve approx. 1.1 as long as 5th

seg. of hind tarsus. Front wing approx. 7.8 mm, with 2nd recurrent vein weak-

ly sloping.

Color: Head, base, apex of mandible, thorax, abdomen, coxae, hind tro-

chanter (except apical margins) black. Median area of mandible ferruginous.

Tegula, extreme bases of tibiae, external surface of foretibia, external surface

of midtibia (except apex), externo-median surface of hind tibia, basal 0.4 of

-hind basitarsus, whitish. Maxillary palpus with apical 4 segments, labial palpus

with apical segment, fore, middle trochanters (except ventral surfaces of basal

segments which are infuscated), apical 0.5 of apical seg. of hind trochanter,

femora, internal surfaces of all tibiae (except bases and apex of hind), apex

of midtibia (externally), ferruginous. Hind tibia with weak sub-basal ferrugi-

nous band, distinct apical brown band. Hind basitarsus with apical 0.6 brown.

Male: Unknown.

Holotype: Female, USSR. Volgograd, Sarepta, coll., Bekker, date? De-

posited in Academy of Sciences, Leningrad.

Etymology: The species is named from the Latin noun 'cribum' meaning

seive, in reference to the densely punctate propodeal trough.

Host and Biology: Unknown.

174 Contrib. Amer. Ent. Inst., vol. 20, 1983

ACKNOWLEDGMENTS

I thank Dr. D. R. Kasparyan (Academy of Sciences, Leningrad) for the

loan of the holotypes of brunipalpus and cribus. I also thank Mr. Roy

Danielsson (Museum of Zoology and Entomology, Lund) for the loan of the

lectotype of Omorguspicticrus Thom. and Mr. Erich Diller (Zoologische

Staatssammlung, Muenchen) for the loan of the holotype of Sesioplex punc-

tulatus Horst. Dr. Klaus Horstmann graciously examined the holotypes of

brunipalpus and cribus before the manuscript was submitted for publication

to be certain that they had not previously been described and placed in other

genera. Special thanks are given to Dr. D. M. Davies (Biology Dept.,

McMaster Univ.) for his financial support of this study (NSERC operating grant).

REFERENCES

Horstmann, K. 1977. Bemerkungen zur Systematik einiger Gattungen der

Campopleginae, II, (Hymenoptera: Ichneumonidae). Mitt. Munch. Ent. Ges.

67: 65-83.

Pielou, E. C. 1979. Biogeography. John Wiley & Sons, New York, Chichester,

Brisbane, Toronto. 35] pp.

Sanborne, M. 1981. A revision of the genus Sesioplex (Hymenoptera:

Ichneumonidae) of North America. Can. Ent. 113: 623-629.

Townes, H. K. 1969. Genera of Ichneumonidae, Part 1. Mem. Am. Ent. Inst.

11. Ann Arbor, Michigan. 300 pp.

Viereck, H. L. 1912. Contribution to our knowledge of bees and ichenumon-

flies, including the descriptions of twenty-one new genera and fifty-seven

new species of ichneumon-flies. Proc. U.S. Natl. Mus. 42: 644.

Wiley, E. O. 1981. Phylogenetics. The theory and practice of phylogenetic

systematics. Wiley-Interscience. New York. 439 pp.

Sanborne: Sesioplex (Ichneumonidae) 175

Figs. 1-8. Sesioplex brunipalpus, n. sp.: 1, Frontal view of head.

2, Dorsal view of head. 3, Propodeum. 4, Dorsal view of first abdominal

tergite. 5, Lateral view of first abdominal tergite. 6, Hind tarsal claw.

7, Hind leg. 8, Ovipositor. Scale = 1.0 mm.

1983

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A REVIEW OF THE WORLD SPECIES OF DYSPETES

(HYMENOPTERA : ICHNEUMONIDAB)

Virendra Gupta

Center for Parasitic Hymenoptera

Department of Entomology & Nematology

University of Florida, Gainesville, FL. 326ll

Dyspetes Foerster is a small genus of the tryphonine ichneumonids belong -

ing to the tribe Tryphonini. Only three species are known so far:

D. arrogator Heinrich (= praerogator Gravenhorst = fracticeps Townes) from

Eurasia, D. orientalis Kasparyan from Eastern Palearctic Region and D.

rufus (Provancher) from Canada. In the present paper three new species,

townesi, flavus and indicus, are described from Japan, Burma and India,

respectively. The genus is recorded from the Oriental Region for the first

time.

The hosts of only D. arrogator are known. They are larvae of the fami-

ly Tenthredinidae.

Genus DYSPETES

Dyspetes Foerster, 1869. Verh. Naturh. Ver. Rheinlande, 25: 201.

Type: (Tryphon praerogator of Gravenhorst, not Linnaeus) = arrogator

Heinrich. Included by Thomson, 1883.

Dyspetus Thomson, 1883. Opuscula Entomologica, 9: 899. Emendation.

Taxonomy: Townes and Townes, 1950: 422. Perkins, 1962: 419. Townes,

Momoi, and Townes, 1965: 105. Townes, 1969: 176. Kasparyan, 1981: 130.

Moderately large, slender species, with slender antennae and long wings.

Body largely closely punctate. Clypeus broad, with a median transverse crest,

its base and apical margin each with a row of long hairs. Mandible moderate-

ly long. Malar space 0.5 to 1.0 as long as the basal width of mandible. Eyes

slightly to moderately diverging ventrally towards clypeus. Occipital carina

prominent, ridge-like, strongly convex to angulate medially above. Sometimes

occiput notched at this point. Posterolateral corner of pronotum not thicken-

ed or projecting. Subtegular ridge sharp and high. Tegula convex. Propodeum

various, strongly distinctly carinated (as in rufus) to almost without carinae

but for the apical transverse carina. Areola usually defined by a smoother

area when median longitudinal carinae indistinct or weak. Areola and basal

area confluent but with a constriction at their junction. Propodeum punctate

except in species with fully areolated propodeum. Areolet rhombic, wider

than high, with a very short petiole above, receiving second recurrent vein near

its middle or a little basad or distad of the middle. Second recurrent vein

almost straight, with two well separated bullae. Nervellus intercepted usually

at its lower 0.35 to 0.4, but in one species a little above the middle. Tarsal

claws pectinate in their basal 0.3. Median dorsal carinae of first tergite rep-

resented by low rounded ridges. Dorsolateral carina of first tergite sharp to

the level of spiracle, beyond which it is obsolete. Second tergite punctate,

with postmedian and sublateral impressions that bound a median swollen area.

Ovipositor sheath flat, decurved, flexible near the base, about 0.45 as long as

the apical depth of abdomen. Ovipositor decurved, about as long as the apical

depth of abdomen. Figure 1 depictsthe generic characters. Figures 2-11 show

other characters of the genus and the included species.

177

178 Contrib. Amer. Ent. Inst., vol. 20, 1983

Fig. 1. Dyspetes arrogator Heinrich, female, generic characters.

(After Kasparyan)

V. Gupta: Dyspetes (Ichneumonidae) 179

KEY TO THE WORLD SPECIES

Propodeum fully areolated. Carinae strong. Costula strong. Propodeum

with scattered punctures or rugosities, subpolished. Occiput with a

shallow indentation in the occipital carina. Ovipositor weakly arched

downwards. Body wholly rufous with some fuscous marks, especially

in the male. Neatctic:-{(Canada) . <6. e . 6. rufus (Provancher)

Propodeum not fully areolated. Costula absent (except rarely in the

males). Often only the apical transverse carina seen or weak median

longitudinal carinae may be present bounding a smoother areola. Pro-

podeum punctate. Occiput deeply notched or not so. Color black or

yellow (one amecies from Butmal.¢7s) 1... oe Ce ee. Z

Occiput with a deep cleft in the middle of occipital carina. Areola shiny,

bounded by weak low carinae. Ovipositor strongly decurved ..... 3

Occiput without a median cleft. Occipital carina may be strongly angled

in this region, but not forming a cleft. Areola variable: absent to

present. Ovipositor strongly to weakly decurved ............ 4

Scutellum convex, ‘its lateral carina confined to its base. Frons without

a median carina, or sometimes with an incomplete one. Mesopleurum

and metapleurum with moderately dense punctures, with interspaces

more than the diameter of the punctures. Claspers rounded and nar-

rower apically. Female subgenital plate sparsely hairy at apex. Hind

leg color variable: wholly black to wholly brownish-yellow. Fore and

middle coxae usually brownish-yellow. Eurasia... . .. ./s ue ws

i ei ke hk a be PMR es use ae lee Aa eee ae ae eg ae a 1. arrogator Heinrich

Scutellum flat, its lateral carina reaching almost to the apex and giving

scutellum an angulate appearance. Frons with a median carina. Meso-

pleurumand metapleurum closely punctate, punctures minute to small,

but dense and close together. Claspers broad, truncate apically. Fe-

male subgenital plate densely hairy at its apex. All coxae black. Hind

leg Blaeic.) japanse dg. CMe. a ee eae 2. townesi ; neep.

Body wholly yellowish-brown with black lines on thorax. Frons with a

shallow median groove and strans-striations laterally. Propodeum wholly

punctate. Areola usually not clearly demarcated glee the rest of pro-

POURURD Ne Gea ie Ot a Pavisee sn ges ie : aflevus ~ Th SP.

Body wholly black, or abdomen banded with black and ae Frons with-

out any groove or carina, closely punctate or rugulose. Areola de-

marcated by a smoother area, or propodeum uniformly punctate... 5

Second and the following tergites yellow, with broad black marks in the

middle (extent variable, sometimes less yellow on tergites). Median

longitudinal carinae of propodeum usually distinct, demarcating a some-

what smoother areola. Frons punctate. Mesopleural punctures not

coarse, interspaces shiny and at least as much as the diameter of the

punctures. Ovipositor weakly decurved. Second recurrent vein emitted

from the middle of the areolet. Japan, Russia (Far East) .......

ee eS Me nT PIR ROR 2 ie MO ay ee a ae 4. orientalis Kasparyan

Abdomen wholly black. Propodeum without distinct areola or median

longitudinal carinae, uniformly densely punctate. In males, however,

weak carinae often seen. Frons rugolose. Mesopleurum closely

180 Contrib. Amer. Ent. Inst., vol. 20, 1983

6, ee he Ss

. pag eis

Figs. 2-8. Dyspetes arogator Heinrich : 2, Head, front view. 3,

Propodeum and first tergite. 4, First and second tergites. 5, Male

genitalia, ventral view. 6, Syntergites of male genitalia. 7, Subgenital

plate. 8, Vertex. Fig. 9, D. rufus (Provancher), tip of female abdomen.

(After Kasparyan and Townes)

V. Gupta: Dyspetes (lchneumonidae) 181

punctate, without smooth or shiny interspaces. Ovipositor strongly de-

curved. Second recurrent vein emitted from beyond the middle of the

areniet..jdpslialt pig Rak io Oreis 2 Un wot +. tindious, tt. Sp.

1. Dyspetes arrogator Heinrich (Figs. 1-8)

Tryphon praerogator Gravenhorst, 1829. Ichneumonologia Europaea, 2: 127.

Misdetermination of Ichneumon praerogator Linnaeus, 1758. Syn. by

Townes, Momoi & Townes, 1965: 105.

Dyspetes arrogator Heinrich, 1949. Mitt. Muenchen Ent. Gesell., 35-39:

107. F, M. Type M, Germany: "Haag a.d. Amper. Obb., May 1948"

(Townes).

Dyspetes fracticeps Townes and Townes, 1950. Ann. Ent. Soc. America,

42(3-4): 421. New name for praerogator Gravenhorst.

Taxonomy: Kasparyan, 1973: 100. Hinz, 1976: 100. Kasparyan, 1981: 131.

This European species is apparently widespread in Eurasia. It exhibits dif-

ferent color forms, showing differences in the color of hind leg, antenna, etc.,

which are probably associated with different seasons or host associations. Hinz

(1976) recognized two distinct subspecies within arrogator: (a) Dyspetes

arrogator arrogator Heinrich, parasitizing Aglaostigma aucupariae (Tenthredi-

nidae) and appearing in early summer (May) in Germany, and (b) D.

arrogator fracticeps Townes and Townes (= praerogator Gravenhorst), para-

sitizing Tenthredo and Tenthredopsis species (Tenthredenidae) and appearing

from June to August. Evidences on whether these differences are seasonal

or whether the two taxa are also geographically isolated are inconclusive at

the moment, as I find both color forms in collections made during July-August

in Europe. Kasparyan (1981) considers the two forms as different generations

(spring and summer-autumn) of the same species. The following description

takes into consideration the variability within the species.

Male and Female: Face coarsely punctate, depressed near base of clypeus.

Basal half of clypeus with distinct separated punctures. Frons punctate, with

a faint incomplete median carina, which is often indistinct. Occipital carina

strong, strongly angled medially above, and making a A -shaped cleft in the

occiput. Interocellar distance about equal to ocellocular distance in female

and about 0.8 in male. Malar space 0.75 (F) and 1.0 (M) the basal width of

mandible. Mesoscutum with minute shallow punctures. Scutellum convex, in-

distinctly punctate, its lateral carina confined to its base. Mesopleurum and

metapleurum with moderately dense punctures, with interspaces shiny and equal

to or more than the diameter of the punctures. Punctures a little closer on

metapleurum. Propodeum punctate, smoother in the region of areola. Areola

incompletely formed. Costula absent, or partly faintly represented, with median

longitudinal carinae angled in the region of costula. Apical transverse carina

the strongest of all propodeal carinae. Nervulus interstitial or a little distad

of basal vein. Second recurrent vein joining areolet in its middle or a little

apicad of the middle. Nervellus intercepted at lower 0.35 to 0.4. Basal

three tergites finely punctate. Rest of the tergites mat. Ovipositor strongly

decurved. Subgenital plate folded medially and with sparse hairs at its pointed

apex. Male claspers somewhat narrowed apically and obliquely truncate.

182 Contrib. Amer. Ent. Inst., vol. 20, 1983

Color: Female: Black. Antenna brown to yellowish-brown. Fore and mid-

dle legs yellowish-brown. Hind coxa black to brown. Rest of hind leg

yellowish-brown with apices of femur, tibia and whole of tarsus, blackish-

brown. Tegula yellowish-brown. Wings with brownish tinge.

Male: Antenna black to brown. Hind leg largely blackish. Hind coxa

black. Fore and middle coxae brown to brownish-black. Color of hind femur

variable from black to yellowish~brown. Fore and middle legs usually yellowish-

brown. The color variations are tabulated below:

Color variations in specimens examined from various localities and months.

Germany May Male Antenna black. Hind leg black. Fore and

middle coxae brownish-black. Legs other-

wise yellowish-brown.

Germany May Female Antenna and all legs brownish-yellow.

Germany July Male Antenna black. Fore and middle legs and

: hind femur yellowish-brown. Hind coxa

black. Fore and middle coxae brown.

Germany July Female Antenna and legs reddish-brown. Hind

coxa black.

Austria July Male Antenna brown. Hind coxa blackish. Hind

femur yellowish-brown. Fore and middle

coxae yellowish-brown.

Italy July Female Antenna yellowish-brown. All legs

yellowish-brown. Hind coxa brown.

England Sept. Male Antenna black. Hind coxa black. Fore

and middle coxae and hind femur brown.

England Sept. Female Antenna brown. All legs yellowish-brown,

with hind coxa black.

Venlo Sept. Female Antenna yellowish-brown. All legs

yellowish-brown. Hind coxa brown.

Length: 7-12 mm. Fore wing 7-12 mm.

Specimens: Germany: "Haag, Amper. Ober Bayern", | M (paratype),

24. V. 1948, G. Heinrich (Townes). Berchtesgaden, Jennenmassiv, 1200 m, 1

M, 1F, 10. VII. 1947 and 22. VII. 1947, G. Heinrich (Townes). Austria: Tirol,

Scheffan, 800 m., 1 M, 25. VII. 1958, D. Townes. Italy: Franzenfeste, 1000 m,

1 F, 8 VII. 1958, D. Townes. Venlo, 1 F, 17. IX. 1942, Teunissen. Denmark:

Copenhagen, 4 M, 1 F, 27. VIII. 1958, D. Townes. England: Oxford, Bagley

Woods, 5 M, 10 F, 25. IX. 1960 and IX. 1980, H. Townes and D. Owen.

Ireland, Wicklow Co., 1 M, 1 F, 22. VII. 1940, B. P. Beirne, Dublin Co., | M,

26. VIII. 1942, A. W. Steltox (All Townes).

Hosts (vide Hinz 1961): Tenthredo livida, T. rubricoxis, Tenthredopsis

carbonaria, T. excisa, T. nassata, and Aglaostigma aucupariae.

Distribution: Europe, USSR.

V. Gupta: Dyspetes (Ichneumonidae) 183

Uchida (1930) and others (see Townes, Momoi and Townes 1965: 106) re-

ported D. praerogator from Japan. According to Townes et al. (1965) these

records were erroneous. The specimens from Japan examined in the Townes

Collection represent two species: D. townesi, n. sp., and D. orientalis

Kasparyan.

Morley (1913) recorded this species from Darjeeling, India by a single fe-

male. This specimen could not be traced, nor any other specimen of it col-

lected so far from India. Townes, Townes and Gupta (1961: 106) placed it as

Dyspetes sp-

2. Dyspetes townesi, n. sp. (Fig. 10)

Male and Female: Face coarsely punctate. Clypeus punctate in basal half.

Malar space 0.75 the basal width of mandible. Frons punctate, with a median

carina. Occipital carina strongly angled medially above, forming a median A -

shaped notch. Interocellar distance 0.5 -0.6 (female) to 0.8 (male) the ocel-

locular distance. Mesoscutum with minute shallow punctures. Scutellum flat

dorsally, with scattered punctures, shiny in between, its lateral carina extending

to the apex of scutellum and giving it an angled appearance. Mesopleurum and

metapleurumclosely punctate, punctures smaller but close together. Propodeum

punctate. Areola smoother and bounded by weak median carinae. Costula ab-

sent but median carinae angled in the region of costula. Apical transverse

carina of propodeum distinct and circularly arched. Nervulus a little distad

of basal vein. Second recurrent vein in the middle of areolet. Nervellus inter-

cepted at lower 0.35-0.4. Abdominal tergites 1 and 2 finely punctate, 3

granulose, and 4 onwards mat. Abdomen sometimes a little smoother in the

male. Ovipositor strongly decurved. Female subgenital plate weakly folded

medially and with dense hairs at its pointed apex. Male claspers broad and

truncate apically.

Black. Flagellum brownish apically. Clypeus reddish-brown apically. Man-

dible yellowish-brown. Tegula brown. All coxae black. Fore and middle legs

yellowish-brown, with middle femur blackish-brown. Hind leg wholly black.

Often fore and middle legs brownish-black and flagellum blackish in the male.

Sometimes black color of body replaced by dark blackish-brown.

Length: 7-l2 mm. Fore wing 8-13 mm.

Holotype: Female, Japan: Kamikochi, 31. VIII. 1954, Townes family (Townes).

Paratypes: 62 M, 24 F, Japan: Kamikochi, 23-31. VII. 1954, Townes family

(Townes).

Distribution: Japan: Honshu Island.

This species and/or D. orientalis have apparently been reported in the

Japanese literature as D. praerogator. Townes et al. (1965: 106) considered

these as misdeterminations and reported them as Dyspetes spp. from Japan,

Kamchatka and Russia. Specimens from Japan in the Townes Collection repre-

sent both D. townesi and D. orientalis.

3. Dyspetes flavus, n. sp. (Fig. 11)

Male: Face punctate. Clypeus a little rugoso-punctate basally. Malar

space 0.8 the basal width of mandible. Frons with a narrow median groove,

trans-striate to striato-punctate laterally. Interocellar distance 0.7 to 0.8 the

184 Contrib. Amer. Ent. Inst., vol. 20, 1983

AREA

A ‘

Bg |

; _

Figs. 10-11. 10, ‘Dyspetes townesi, vertex showing notched

occipital carina. 11, D. flavus, vertex, showing rounded

occipital carina.

V. Gupta: Dyspetes (Ichneumonidae) 185

ocellocular distance. Occipital carina roundly arched, without any angulation

or indentation in the middle. Mesoscutum finely punctate. Scutellum flat

dorsally and punctate, its lateral carina giving a weak angulation to the

scutellum on the sides. Mesopleurumand metapleuruma little coarsely punc-

tate, with punctures coalescing in the middle. Propodeum punctate. Median

longitudinal carinae faint to absent. Areola absent or not properly formed.

Nervulus interstitial. Second recurrent vein emitted from the center of areol-

et. Nervellus intercepted at upper 0.35 to 0.4. Abdominal tergites | to 3

punctate, rest granulose to mat. Male claspers broadly rounded apically.

Yellowish-brown. The following areas with blackish marks: Mesoscutal

lobes, base of scutellumyupper "neck" region of pronotum, area along prepectal

carina to subtegular ridge, sternaulus, mesosternum, mesepimeron, along sub-

metapleural carina, basolateral aspects of tergite 1, and sides of abdomen.

Apical abdominal tergites yellowish-brown. Hind tibia & apex of femur light

brown.

Female: Unknown.

Length: |i mm. Fore wing ll mm.

Holotype: Male, Burma: Mt. Victoria, 2600 m, V. 1938, G. Heinrich

(Townes).

Paratypes: 2 M, same data as the type (Townes).

Distribution: Burma.

4. Dyspetes orientalis Kasparyan

Dyspetes orientalis Kasparyan, 1976. Trudy Biologo-Pochvennogo Instituta,

Novaya Seriya, 43(146): 108. M, F. des. Type: F, USSR: Sernovodsk

(Leningrad). Kuril Islands. China. USSR.

Male and female: Face punctate, tending to be rugose at places. Clypeus

with scattered punctures in the basal half. Malar space 0.4 to 0.5 the basal

width of mandible. Frons punctate, without a carina. Interocellar distance

about 0.6 to 0.75 the ocellocular distance. Occipital carina a little strongly

curved or broadly angulate medially, but not forming any cleft. Mesoscutum

finely punctate. Mesopleurum and metapleurum punctate, interspaces shiny and

at least as much as the diameter of the punctures. Propodeum punctate, with

its central area smooth or subpolished and with irregular punctures. Areola

usually demarcated by weak to stronger median carinae, which are often angled

in the region of costula, though costula absent. Apical transverse carina of

propodeum strong and arched. Nervulus distad of basal vein. Second recurrent

vein emitted from the middle of areolet. Nervellus intercepted in lower 0.35

to 0.4. Abdominal tergites 1 to 2 punctate, 3 with scattered punctures to mat,

and rest of the tergites mat. Ovipositor weakly decurved. Female subgenital

plate folded medially, with moderate hairs at its pointed apex. Male claspers

slightly tapered and truncate apically.

Black. Clypeus reddish-brown. Mandible yellow. Antenna blackish in the

male and yellowish-brown in the female. Tegula yellow. Fore and middle

legs yellowish-brown. Coxae often brownish. Hind coxa and femur black, its

trochanters, tibia and tarsus yellow with apical 0.3 to 0.5 of tibia black to

blackish-brown. Abdomen black, with tergites 2 to 4 often yellowish-brown

186 Contrib. Amer. Ent. Inst., vol. 20, 1983

and with a black central mark of various shape. Fourth and the following ter-

gites largely black to largely yellowish-brown. Abdomen has a banded ap-

pearance. | |

Length: 7-l1 mm. Fore wing 7-12 mm.

Specimens: Russia: 1 M, 1 F (paratypes), 16-20. VIII. 1973, Kasparyan

Coll. (Townes). Japan: Kamikochi, 91 M, 64 F, 22-26. VII. 1954, Townes

family (Townes). Mt. Norikura, 1500-2000 m, 1 M, 1 F, 30-31. VII. 1954, Townes

family (Townes). Nagano Pref., 2 M, 2. VIII. 1931, Kaku Sato (Townes). Nikko,

1 F, 18. VII. 1931, J. L. Gressitt (Townes).

Distribution: Japan: Honshu Island, USSR, Kamchatka.

A male from India: Kumaon Hills: Dwali, 2743 m, 8. X. 1973 (Gupta

Collection) comes close to D. orientalis, but has the occipital carina a little

more strongly angulate medially above, tegula blackish-brown, middle coxa and

hind leg wholly black, and abdominal tergites black with narrow apical yellow

bands on tergites 2-4. It may represent an Indian subspecies of D.

orientalis. |

5. Dyspetes indicus, n. sp.

Male and female: Face punctate to rugoso-punctate. Clypeus flatter, some-

what rugulose to ruguloso-punctate. Malar space 0.7 to 0.85 the basal width

of mandible. Frons rugulose to ruguloso-punctate, without any carina, a little

smoother in the middle line. Interocellar distance 0.6 to 0.75 the ocellocular

distance. Occipital carina strong, uniformly arched and rounded medially above.

Mesoscutum finely punctate. Scutellum convex, closely finely punctate, its

lateral carina confined to its base. Side of thorax closely punctate, punctures

often coalescing and without smooth interspaces. Propodeum closely punctate,

as is metapleurum a little rugulose medially, without a distinct areola or median

longitudinal carinae. Apical transverse carina of propodeum also broadly erased

medially. Nervulus distad of basal vein. Second recurrent a little beyond the

middle of areolet. Nervellus intercepted at lower 0.35 to 0.4. Tergites 1, 2

and 3 finely closely punctate, the rest mat. Ovipositor strongly decurved.

Female subgenital plate folded medially, its apex with sparse hairs. Male

claspers broadly rounded apically.

Black. Clypeus, mandible, flagellum, legs, and tegula yellowish brown to

reddish-brown. Flagellum basally often black. Face sometimes with a reddish-

brown median mark. Hind coxa and tibia sometimes black marked.

Length: 10-12 mm. Fore wing 9-ll mm.

Holotype: F, India: Himachal Pradesh: Kalatop, 2438 m, 4. IX. 1971, JD

100 (Gupta).

Allotype: Male, India: Ahla, near Kalatop, 2286 m, 2. IX. 1971, JD 92,

(Gupta).

Paratypes: 2 M,1F, Uttar Pradesh: Valley of Flowers, Garhwal Himalaya,

VI. 1978, V. K. Gupta (Gupta). Nainital, 1939 m, in Kumaon Hills, | F, 7. IX.

1972, Gupta. Himachal Pradesh: Ahla, 2286 m, 2 M, 1 F, 16. IX. 1971, JD 92,

JD 139. Kalatop, 2438 m, 7 M, 9 F, 31. VIII. to 17. IX. 1971. Simla Hills:

Kufri, 8200 ft, 4 F, 12. X. 1966, J 180. Narkanda, 9000 ft, 2 F, 6. X. 1962,

Gupta. (All Gupta).

V. Gupta: Dyspetes (Ichneumonidae) 187

Distribution: India: Uttar Pradesh and Himachal Pradesh Hills.

6. Dyspetes rufus Provancher (Fig.9)

Exetastes rufus Provancher, 1874. Nat. Canad., 6: 78. M, F. des.

Lectotype (designated by Gahan and Rohwer, 1917): F, Quebec (Ste

Foy) (Quebec). |

Male and female: Face moderately closely and strongly punctate. Vertex

and ocellar area with well separated punctures, shiny in between punctures.

Frons without a median carina; there may be a shallow groove instead of a

carina. Occipital carina with a distinct angulation medially above, but occiput

not notched at this point. Vertex depressed near the angulation in occipital

carina. Mesoscutum shallowly punctate. Scutellum convex, its lateral carina

extending up to 0.75 its length. Mesopleurum and metaplurum moderately

coarsely punctate, punctures separated from each other by about 1.3 times

their diameter. Propodeum fully areolated, its carinae sharp and high, with

indistinct punctures or shallow rugosities. Costula sharp and distinct. Areolet

receiving second recurrent vein a little basad of its middle. Nervulus a lit-

tle distad of basal vein. Nervellus intercepted at its lower 0.4. Second ter-

gite subpolished with weaker punctures separated by about 1.5 their diameter.

Ovipositor sheath rather flat. Ovipositor not sharply decurved, only a little

arched downwards.

Ferrugineus. Apical 0.4 of flagellum, fore and middle tibiae and all tarsi,

fulvo-flavous. Mesosternum, hind femur, apical 0.25 of hind tibia, and second

and third tergites faintly, often with blackish tinge. Male comparatively darker

with fuscous marks on mesopleurum, mesoscutum, metapleurum, hind coxa, hind

femur more fully, dorsum of propodeum and all abdominal tergites. Wings

yellowish-hyaline.

Length: 9-10 mm. Fore wing 8.5-9.0 mm.

Specimens: Canada: Finland, Ontario. | M, 21. VII. 1960, S. M. Clark.

North Branch, Ontario, | M, 23. VII. 1960, S. M. Clark. One Sided Lake,

Ontario, 2 F, 24. VII. 1960, S. M. Clark and 1. VIII. 1960, Kelton and Whitney.

Beause Joure, Manitoba, 1 F, 20. VIII. 1953. Brooks-Kelton (All Ottawa).

North Branch, Ontario, | F, 23. VI. 1960, S. M. Clark (Townes).

Distribution: Canada: Ontario, Manitoba and Quebec.

REFERENCES

Heinrich, G. 1949. Neue und interessante Schlupfwespen aus Bayern (Hym.).

Mitt. Munchen Ent. Gesell. 35-39: 101-127.

Hinz, R. 1961. Uber Blattwespenparasiten (Hym. und Dipt.). Mitt. Schweiz.

Ent. Gesell. 34(1): 1-29.

Hinz, R. 1976. Zur Systematik und Oekologie der Ichneumoniden V. (Hymen-

optera). Deutsch. Ent. Ztschr. 23(1-3): 99-105.

Kasparyan, D. R. 1973. Najesdniki-ichneumonidi (Ichneumonidae) Podsemejstwo

Tryphoninae. Triba Tryphonini. Fauna SSSR, Nassekomije

perepontschatokrylije 3(1): 1-319. (In Russian).

188 Contrib. Amer. Ent. Inst., vol. 20, 1983

Kasparyan, D. R. 1976. Trudy Biologo-Pochvennogo Instituta, Novaya Seriya

43(146): 107-120. (In Russian). English translation: New species of

Ichneumon flies of Tribe Tryphonini (Hymenoptera, Ichneumonidae) from

the southern part of the Far East. Appendix I to Fauna of the USSR,

1981.

Kasparyan, D. R. 1981. Fauna of the USSR. Hymenoptera Vol. III, No. 1,

Ichneumonidae (Subfamily Tryphoninae), tribe Tryphonini. Amerind

Publ. Co., New Delhi, pp. 1-388. Appendix I: pp. 391-400.

Morley, C. 1913. Fauna of British India, Hymenoptera, Vol. III. Ichneumon-

idae. 531 pp. Taylor and Francis, London. Reprinted by Today &

Tomorrow's Printers and Publishers, New Delhi.

Townes, H. K. and Townes, M. C. 1950. A _ revision of the genera and of

the American species of Tryphoninae, Parts I and II]. Ann. Ent. Soc.

Amer. 42(3-4): 321-447.

Townes, H., Townes, M., and Gupta, V. K. 1961. A catalog and reclassifica-

tion of Indo-Australian Ichneumonidae. Mem. Amer. Ent. Inst. 1: 1-501.

(Dyspetes, pp. 105-106).

Townes, H. Momoi, S., and Townes, M. 1965. A catalog and reclassification

of Eastern Palearctic Ichneumonidae. Mem. Amer. Ent. Inst. 5: 1-661.

(Dyspetes, pp. 105-106).

Townes, H. 1969. The genera of Ichneumonidae, part I. Mem. Amer. Ent.

Inst. Il: 1-300.

A NEW SPECIES OF ASTICHUS PARASITIC ON A

CIID BEETLE FROM BORNEO

(HYMENOPTERA : CHALCIDOIDEA : EULOPHIDAE)

Tetsusaburo Tachikawa

Entomological Laboratory, College of Agriculture

Ehime University, Matsuyama, Japan

Through the courtesy of Mr. Masahiro Sakai, I recently had a chance to

examine two female specimens which were reared from Cis sp. (Coleoptera:

Ciidae) in Borneo Is., Malaysia, and I take this opportunity of describing a new

species of the genus Astichus Foerster for the first time from Borneo.

Astichus townesi, n. sp.

The present new species is so peculiar as to be unable to put through

the key to the female Astichus species given by Hedqvist (1969). The female

of this species may be characterized by the distinct maculation and shape of

the fore wings and the coloration of the funicle.

Female: Head black and metallic greenish blue except for the mouth

parts which are yellowish borwn; pronotum, mesoscutum and axillae black with

metallic greenish blue; scutellum, propodeum and abdomen black with vivid,

metallic purple lustre. All funicle segments yellowish white; scape (except

for the basal and apical whitish portions), pedicel and club blackish. All coxae

and trochanters blackish; all femora, tibiae and tarsi yellowish white except

for the following dark brownish parts: basal halves of femora and tibiae,

apical portions of tibiae and 2nd segments of tarsi. Maculation of fore and

hind wings shown as Fig. l, A and C.

Head, seen from above, about 1.7 times as wide as frontovertex; fronto-

vertex concave medio-anteriorly; ocelli in an obtuse-angled triangle; a

transverse stripe* in front of median ocellus white. Antennae inserted on the

lower eye level; scape a little expanded on apical half, exclusive of the radicle,

about as long as club; pedicel slightly longer than the Ist funicular segment;

two ring segments very tiny; funicle 4-segmented; the Ist funicular segment

much shorter than the 2nd; the 2nd to 4th funicular segments pedunculate and

subequal in length.

Pronotum short; notaulices (= parapsidal grooves) deep and complete,

scapulae (= side lobes) convex; scutellum about as long as wide. Gaster elon-

gate, a little longer than head and thorax taken together; cercal plates situated

near the top of abdomen; ovipositor sheath exserted, about one-third the length

of gaster.

Length: 1.2 mm.

Male: Unknown.

Host: Cis sp. (Coleoptera: Ciidae) feeding on a fungus.

Holotype: F, Polong, Sabah, Borneo Is., Malaysia, 18. V. 1980, M. Sakai

leg., ex Cis sp. Paratype: 1 F, the same data as holotype. The type~series

A. mi {uatakes Tachikawa, 1977 and A. sakaii Tachikawa, 1977, this whitish

sttipe is lacking.

189

190 Contrib. Amer. Ent. Inst., vol. 20, 1983

is deposited in the Entomological Laboratory, Ehime University. This new

species is named in honor of Dr. H. K. Townes.

I thank Mr. Masahiro Sakai (School of Medicine, Ehime University) for

gift of the specimens, and Assoc. Prof. Mutsuo Miyatake (College of Agricul-

ture, Ehime University) for identity of the host ciid beetle.

REFERENCE

Hedqvist, K.-J. 1969. Notes on the genus Astichus Foerst. and description

of new species (Hym., Chalcidoidea, Eulophidae, Euderinae). Entomol.

Ts. 90 (3-4): 166-173.

Fig. 1. Astichus townesi, n. sp., female: A, fore wing.

B, mandible. C, hind wing. D. antenna. (Photo: Tachikawa).

A NEW GENUS OF ENDASEINE ICHNEUMONIDAE

FROM AUSTRALIA (HYMENOPTERA)

I. D. Gauld

Department of Entomology, British Museum (Natural History)

London, U.K.

G. A. Holloway

Department of Entomology, Australian Museum

Sydney, Australia

Abstract

A new endaseine genus, Tryonocryptus, and three new species, T.

nigridorsalis (the type-species), T. gigas and T. amicus, are described from

Australia. Meringops Townes, a genus previously known only from South

America, is newly recorded from Australia and a key is given to separate

these two genera. The systematic and zoogeographical relationships of the

Australian endaseines are discussed.

INTRODUCTION

The subtribe Endaseina is one of the most distinctive groups of

Phygadeuontinae (= Gelinae). They are rather stout ichneumonids with short

spinose tibiae and, although little is known of their biology, their stout legs

suggest they search for hosts amongst leaf litter and in the soil. A few

species have been reared from cocoons of Lepidoptera and Symphta and one

Nearctic species parasitizes dytiscid pupae. The majority of endaseines are

restricted to cooler humid areas. Most species occur in the north temperate

region and montane South America, though a few species are known from

mountains in South East Asia (Townes 1970).

A recent study of Australian ichneumonids has revealed that two enda-

seine genera occur on the continent. One, Meringops, was previously only known

from Chile, whilst the other, herein newly described, is strikingly different from

all known endaseines.

Subtribe ENDASEINA

This subtribe may be recognized with Townes' (1970) key. Some con-

fusion may occur with the subtribe Mastrina but the two are easily separated

by the shape of the scape. That of endaseines is almost transversely trun-

cate, whereas that of mastrines is very obliquely truncate. Mastrines also

have more slender legs, the tibiae are not densely spinose and the central en-

tennal segments of females are usually elongate, not quadrate or transverse

as in most endaseines.

KEY TO ENDASEINE GENERA OCCURRING IN AUSTRALIA

Mandibles slightly tapered with lower tooth slightly shorter than the upper;

clypeus with apex simply arcuate, without a median tooth or tubercle;

upper valve of ovipositor elongately tapered to an acute point .....

aaah gl goin e aM EN rae GT RIC Nia Naleg ae cccigear geal wy Vidar tai Ma a eR Meringops Townes

192 Contrib. Amer. Ent. Inst., vol. 20, 1983

Mandibles distally flared, with lower tooth distinctly the longer; clypeus with

a median apical tooth or tubercle; upper valve of ovipositor bluntly

PU OP BER Oe ees ony Wea ee as Tryonocryptus, n. gen.

Genus MERINGOPS'- Townes

Meringops Townes, 1970. Mem. Amer. Ent. Inst. 12: 80.

Type-species: Meringops minipes Townes, by original designation.

This small genus is known from a single Chilean species. Three further

undescribed species occur in Australia. These species agree well with 'the

generic description given by Townes (1970: 80) except that the fore wing

length should now have a range 4.0 - 6.5 mm.

The Australian species all occur in the cooler, more humid south-

eastern part of the continent. They form a compact species-group and are

clearly closely related to the Chilean species. This type of trans—Antarctic

relationship is fairly uncommon amongst Ichneumonidae, but it is not entirely

unknown. The labenine genera Certonotus, Labium and Labena are virtually

restricted to South America and Australia (Gauld 1983) and the mesostenine

genus Anacis is rich in species in both Chile and south-eastern Australia. It

is thought that these genera dispersed across the Antarctic continent before

the Eocene when the archipelagic connection between Australia and South

America was finally severed as Australia moved northwards. In the Oligocene

and earlier, the South Polar region is thought to have had a warmer climate

than today and there is evidence to suggest the existence of extensive

Nothofagus and southern conifer forests on Antarctica (Rich 1975). It is note-

worthy that even today many of the southern elements in the Australian fauna

are centered in cool temperate forests (Naumann 1982).

Genus TRYONOCRYPTUS, new genus

Type-species: Tryonocryptus nigridorsalis , n. sp. (Fig. 1)

Fore wing length 10-18 mm; head large, body stout; head, alitrunk and

legs (especially of female) densely pubescent. Eye surface with minute in-

conspicuous hairs. Clypeus wide, fairly flat, with a strong median apical tooth

or tubercle, margin blunt. Mandibles stout, distally flared, with lower tooth

conspicuously the longer. Flagellum of female short, stout, with central seg-

ments transverse, that of male more slender, without tyloids; flagellum of both

sexes with a white band. Mesoscutum polished, punctate with notauli vestigial;

scuto-scutellar groove wide, simple; scutellum weakly convex, without lateral

carinae. Mesopleuron with sternaulus short, shallow. Propodeum abruptly

declivous, with anterior and often posterior transverse carinae discernible,

usually close together; median longitudinal carinae absent. Fore wing with

3r-m present enclosing a fairly large pentagonal areolet; 2m-cu with one or

two bullae; postero-distal corner of second discal cell obtuse. Hind wing with

distal abscissa of Cu, present; first abscissa of Cu, shorter than cu-a. Gaster

with tergite 1 slender anteriorly, posteriorly evenly broadened, dorsally flat,

without median lojgitudinal carinae; apex of first sternite just before spiracle.

Tergites 2-4 polished, virtually impunctate, with laterotergites folded under

but not separated by a crease. Female subgenital plate large; ovipositor

Gauld & Holloway: Endaseine Ichneumonidae 193

projecting beyond apex of gaster by 0.7 - 1.6 times length of hind tibia,

slightly decurved and with upper valve deep, apically abruptly and bluntly

rounded.

Tryonocryptus occupies an isolated position within the Endaseina, dif-

fering from all described genera in the form of the mandibles, the medianly

protuberant clypeus and the stout ovipositor. The majority of endaseines share

two apomorphic features ~ a transverse membranous suture at the hind margin

of the mesoscutum and a median longitudinal ridge dividing the scuto-scutellar

groove into two halves. These features are absent in Tryonocryptus though

there is a trace of the suture laterally in some specimens and a slight median

thickening of the scuto-scutellar groove in a few individuals. Of the five

genera without these apomorphies three, Grasseiteles, Charitopes and

Cisaris, are morphologically specialized, but two, Meringops and Medophron,

are both closely related and structurally unspecialized. It is possible that

Tryonocryptus is related to these two genera, but at present no convincing

synapomorphies have been found to confirm this relationship.

KEY TO: SPECIES OF TRYONOCRYPTUS

1. Clypeal margin sinous, with weak lateral lobes and a median truncated

tooth (Fig. 2); mesoscutum black with scutellum ivory marked; oviposi-

tor projecting beyond apex of gaster by 0.7 -~ 0.9 times length of hind

PIM a ska ae a ae oe rae er a a ae ie as lL. Migrigoraalis, Ds sp.

Clypeal margin convex with a large rounded median tubercle (Fig. 3);

mesoscutum and scutellum reddish; ovipositor projecting beyond apex

of gaster by 1.2 or more times length ‘of hind tibia . .. 2.24. 2. 2

2. Ovipositor projecting beyond apex of gaster by 1.5 - 1.6 times length of

hind tibia; mesoscutal punctures obsolescent; large species, fore wing

length 15-18 mm; Victoria/New South Wales ....2. gigas n. sp.

Ovipositor projecting beyond apex of gaster by 1.2 -1.3 times length of

hind tibia; mesoscutal punctures fine but deep; medium sized species,

fore wing length 12~14 mm; Queensland ...... 2, abi cus sn.

1. Tryonocryptus nigridorsalis, n. sp. (Figs. 1-2)

Fore wing length 10-12 mm. Mandibles stout, the lower tooth 2.0 times

the length of the upper; clypeus transverse, with margin sinuate, with weak

lateral lobes (weaker in male than female) and with a truncate median apical

tooth. Flagellum of female short and stout, with 24-26 segments, with a

white band on about segments 12 ~22, those of male more slender with about

30 segments and with white band on segments 13-18. Frons polished, sparsely

punctate, with fewer shorter hairs than face and genae. Mesoscutum polished,

punctate; notauli indistinguishable. Mesopleuron polished, punctate, speculum

without punctures; sternaulus barely discernible, extending less than 0.4 of |

length of pleuron. Propodeum polished, sparsely punctate; anterior transverse

carina present, medially curved forwards; posterior transverse carina present

at least as low lateral keels, when present medially then close to and parallel

with anterior carina. Fore wing with areolet often slightly higher than broad;

2m-cu with two bullae, angled between them and sometimes with a stub of

spurious vein present. Ovipositor projecting beyond apex of gaster by 0.7 -0.9

times length of hind tibia.

194 Centrib. Amer. Ent. Inst.; vol. .20, 1983

Head black; mandibles except teeth, clypeus except apical margin, face,

genae, spot on vertex extending to upper inner orbits and a spot on the tem -

poral orbit, ivory. Pronotum black, near hind corner, ivory; mesoscutum

black, scutellum ivory. Mesopleuron variable, from black to reddish brown,

propodeum similarly colored; metanotum black with postscutellum ivory. Gaster

reddish brown, but with tergite 1 black from base to spiracle. Ovipositor

valves black. Legs reddish brown, hind leg with distal three tarsal segments

white.

Holotype: F, New South Wales: Blue Mts, Mt Tomah, iii. 79 (N.W. Rodd)

(AM). Paratypes: Australian Capital Territory: 1 F, Black Mt, xi. 79 (D.

Colless) (ANIC). New South Wales: 1 M, Blue Mts, Mt Tomah, iii. 79 (N.W.

Rodd) (AM); 1 F, Ben Lomond, ii. 30 (NMV); 1 F, Cabramatta, x. 63 (M. I.

Nikitin) (BMNH); 1 M, Clarence, iii. 79 (n. W. Rodd) (AM); 2 M, Jindabyne,

ii. 69 (A. Neboiss) (NMV). Queensland: 1 M, Brisbane, ix. 12 (H. Hacker) (QM);

1 F, Brisbane, ix. 56 (A. N. Burns) (NMV); 1 M, Mackay (R. E. Turner) (BMNH);

1 F, M'borough, xii. 53 (Barrie) (QUM). Victoria: 1 M, Belgrave, iii. 49 (A.N.

Burns) (NMV); 1 F, Dartmouth Survey, Eight Mile Creek, iv. 53 (NMV); 2 F,

5 M, Fernshaw, i-iii. 55 (A. N. Burns) (NMV); 1 M, Ferntree Gully, xi. 31

(A. N. Burns) (NMV); 1 M, Kallista, iv. 54 (A. N. Burns) (NMV); 2 M, 12 km

SE of Merrijig, xi. 71 (A. Neboiss) (NMV); 1 M, Millgrove, iii. 54 (A. N. Burns)

(NMV); 1M, Mt Hotham, iii. (TC); 1 F, Whisky Flat, i. 46 (A. N. Burns)

(NMV); 1M, Won-Won, ii. 52 (A. Neboiss) (NMV).

2. Tryonocryptus gigas, n. sp. (Fig. 3)

Fore wing length 15-18 mm. Mandibles stout, with lower tooth about 2.0

times as long as the upper; clypeus transverse, margin convex with a rounded

median apical tubercle and no indication of lateral lobes. Flagellum of female

short and stout, with about 28 segments and with white band on segments

13 ~22, those of male more slender, with 31 segments and with white band on

segments 13-19. Frons polished, sparsely punctate, with fewer shorter hairs

than face and genae. Mesoscutum polished, punctures obsolescent, notauli very

weak but indicated almost to posterior margin. Mesopleuron polished, punctate,

speculum smooth; sternaulus vestigial. Propodeum polished, with anterior and

posterior transverse carinae present, the former angled slightly forwards medial-

ly, the latter high laterally, absent centrally or if indicated closely parallel

to anterior carina. Fore wing with areolet broader than high; 2m-cu with one

bulla or two very close. Ovipositor projecting beyond apex of gaster by 1.5 -

1.6 times length of hind tibia.

Head black; mandibles except teeth, clypeus except apical margin, face

except for a median longitudinal area, genae to temporal orbits and inner or-

bits from face to vertex ivory. Antennae, except for white band, black.

Alitrunk and gaster reddish brown, upper hind corner of pronotum ivory. Ovi-

positor valves black. Legs reddish brown, hind leg with distal three tarsal seg-

ments whitish.

Holotype: F, New South Wales: Blue Mts, Mt Tomah, xi. 77 (N. W. Rodd)

(AM). Paratypes: New South Wales: | F, Blue Mts, Mt Bell, xii. 81 (N. W.

Rodd) (AM); | F, 9 miles S of Hampton, ii. 63 (Chinnick and Lewis) (ANIC);

1 F, Jindabyne, ii. 69 (A. Neboiss) (NMV); | M, Tubrabucca, xii. 46 (A. N.

Burns) (NMV). Victoria: | F, Blackburn V., ii. 46 (A. N. Burns) (NMV);

F = female. M = male.

Gauld & Holloway: Endaseine Ichneumonidae 195

6 F, 1 M, Deer Park, ex Entometa apicalis (Walker) (Lep., Lasiocampidae),

ix-x. 47 (A. N. Burns) (NMV); 1 F, Ferntree Gully, iii. 31 (A. N. Burns)

(NMV); 1 M, Kallista, xii. 47 (A. N. Burns) (NMV); 1 F, Nowa Nowa, ii. 52

(A. N. Burns) (NMV); 1 F, Victoria (NMV); 1 M, Victoria (BMNH).

3. Tryonocryptus amicus, n. sp.

Similar to T. gigas except for the following: fore wing length 11 -14

mm; mandibles with lower tooth about 1.5 times as long as the upper; female

flagellum with 24-26 segments, white band on segments 1] ~20, that of male

with about 28 segments, white band on segments 11-19. Mesoscutum with

notauli barely discernible, centrally with fine deep punctures; sternaulus short.

Fore wing with two bullae in 2m-cu. Ovipositor projecting beyond apex of

gaster by 1.2 -1.3 times length of hind tibia.

Coloration as T. gigas except that the facial mark is reddish brown and

indistinct and the ivory mark on the temporal orbit is absent. The entire

alitrunk and gaster is reddish or yellowish brown.

- Holotype: F, Queensland: Bundaberg, Baldwin Swamp, iv. 72 (H. Frauca)

(ANIC). Paratypes: Queensland: 4 M, same data as holotype (ANIC); 1 M,

Biggenden, Coastal Range, v. 74 (H. Frauca) (ANIC); | F, Brisbane, v. ll (QM);

1 F, Brisbane, ix. 20 (H. Hacker) (QM); 1 F, Mt Tambourine, x. 78 (Galloway)

(BMNH); | M, Wybera, xi. 69 (E. C. Dahms) (QM).

ACKNOWLEDGMENTS

We would like to thank Dr. H. Townes for his comments on the place=:

ment of these Australian species. We are grateful to Dr. E. C. Dahms,

Queensland Museum, Brisbane (QM), Dr. I. D. Naumann, Australian National

Insect Collection, Canberra (ANIC), Dr. H. Townes, American Entomological

Insitute, Ann Arbor (TC) and Dr. K. Walker, National Museum of Victoria,

Melbourne (NMV) for the loan of valuable specimens. Part of this work was

undertaken whilst I. D. Gauld was in receipt of an ABRS grant and thanks are

due to the Australian Government for this. Jonathan Carter drew Figure l.

REFERENCES

Gauld, I. D. 1983. The phylogeny of the Labeninae, an ancient southern group

of Ichneumonidae. Syst. Ent. 8:

Naumann, I. D. 1982. Systematics of the Australian Ambositrinae (Hymenop-

tera: Diapriidae), with a synopsis on non-Australian genera of the sub-

family. Aust. J. Zool. Suppl. Ser. 85: 1-239.

Rich, P. V. 1975. Antarctic dispersal routes, wandering continents, and the

origin of Australia's non-passeriform Avifauna. Mem. Natl. Mus. Victoria

36: 63-126.

Townes, H. 1970. Genera of Ichneumonidae. 2. Mem. Amer. Ent. Inst. 12:

[-537,

SEPP gg NR \

5a EN VY, Ae

ee ee oe

1983

= Ee yay ee

~ Se

‘J

meno crepe naman,’

Contrib. Amer. Ent. Inst., vol. 20,

a) a!

196

dorsalis, female, lateral view

nigri

Fig. 1, Tryonocryptus

of the whole insect.

Gauld & Holloway: Endaseine Ichneumonidae 197

Figs. 2-3. Head showing face: 2, Tryonocryptus nigridorsalis,

female. 3, T. gigas, female.

TWO INTERESTING SULCOMESITIUS SPECIES FROM AFRICA

(HYMENOPTERA : BETHYLIDAE)

L. Moczar

Department of Zoology, Hungarian Natural History Museum,

1088 Budapest, Hungary

Abstract

Two Sulcomesitius Méczaér species are recorded from Middle and South

Africa: S. consimilis Méoczar and S. townesianus, n. sp.

1. Sulcomesitius consimilis Moczar, 1970

Speciman: 1 M, Kurger Natl. Park Pretoriushop 21.11.1968, K. V. Krombein

(Paratype no. 255).

Distribution: South Africa: Umtata, Port St. John, Cape Province

(Moczar 1970).

2. Sulcomesitius townesianus, n. sp.

Male: Length 4.6 mm. Black, mandibles, clypeus, antennal joints reddish

brown, lower side lighter, upper side darker; upper side of scape and top of

joints 3-10 more or less with black spots, last 3 joints entirely black; articu-

lating parts of legs dark brown; top of tibiae, tarsal joints, as well as, abdom-

inal segments 3-6 yellowish brown. Wings normal, fore wings only weakly

brownish infuscated with broad lighter band in distal half and outside of cells,

veings brown. Body sparsely covered with short light hairs, antennae with very

short brownish hairs.

Head as long as broad including eyes (28:27) wider and parallel behind

eyes, posterior margin nearly straight with very deep and depressed occipital

margin (Fig. 1), latter with a row of larger punctures; ocelli in a right angle,

POL:001 = 4:6, hind ocelli separated from occiput by the length of OOL (6:6),

outer margins of ocelli with narrow grooves; frontal sulcus not developed; head

coarsely and densely punctured; anterior margin of clypeus semicircularly

rounded and raised as a longitudinal sharp and high keel medially; eye rather

convex, hardly longer than wide (11:10), separated from mandible by hardly

longer of its half length (6:5.5) and separated from occiput by twice the

length of POL. Antennae unusually thick, joints generally only slightly longer

than thick, joint 7 as long as broad, inner sides of joints 3-10 (especially

4-7) curved, outer side straight, outer tips of these joints acute, inner ones

nearly rectangular (Fig. 2), length (and breadth) proportions of joints 1-13 =

9.5/4/: 4/3/: 5.5/4/: 6/5/: -$.5/5/: 6/5.5/: 5/5/: 5.5/4. 573/: 5/3/: 4.5/3/: 4.5/3/:

6/3/. Pronotum slightly longer than half broad in. front (13:23) (including an-

terior corners), anterior corners in front sharply triangularly, protruding owing

to wrinkle emerging in lateral side and reaching corners (Figs 1, 4), lateral

sides of pronotum concave up to middle and diverging backwards (seen from

above), posterior margin only weakly arched, surface very coarsely and deeply

punctured, with larger punctures than vertex, longitudinal furrow deep and nar-

row especially in front (Fig. 1). Mesonotum shagreened, weakly shining with

some smaller punctures, notaulices broad, very deep here sharply cross

wrinkled (Fig. 4) and sharp margined, parapsidal furrow weakly developed,

mesonotum outsides notaulices with larger scattered punctures, longitudinal

furrow broad and deep, present only posteriorly (Fig. 3). Mesonotum well

198

Moczar: Sulcomesitius (Bethylidae) 199

Figs. 1-6. Sulcomesitius townesianus, n. sp. male:

1, head,- pronotum. 2, antenna. 3, mesonotum-propodeum.

4, pronotum-mesonotum. 5, lateral aspect of propodeum.

6, abdomen.

200 Contribs Amer Ents insti, vot. s20; 1983

separated from scutellum by a transverse groove and by a pair of pits at its

base laterally. Scutellum shagreened only basally without median sulcus and

with deeper larger punctures on its whole surface. Dorsal surface of propo-

deum strongly sculptured, and well separated and transversal wrinkled central,

sublateral and lateral areas (Fig. 3), propodeum as long medially as half

breadth of disc transversally (14:14), lateral sides parallel only spines slightly

divergent, lateral spines very acule and conspicuously long, nearly as long as

three-fourth length of propodeum medially (10) (Fig. 3), lateral sides of pro-

podeum with dense larger punctures and with a deep sulcus along hind, as

well as, upper margin, the latter runs up to tip of spine (Fig. 5). Episternum

deeply coarsely and irregularly punctured with a diagonal impression.

Abdomen smooth shining, tergite | polished with scattered distinct punc-

tures medially and laterally, tergite 2 granulated on its one-third part basally,

with distinct, rather deep but more scattered punctures medially (Fig. 6), with

more dense punctures laterally and polished before posterial margin only me-

dially. Last tergites alutaceous basally and finely punctured distally. Sternite

2 deeply and densely punctured.

Holotype: M,Uganda: Ankole Kichwamba, IV. 23-29. 1968, P. J. Spangler

(Washington, USNM Type No. 100681).

This new species together with S. crassiantennis Moczar, 1976 differ from

all other species of the genus Sulcomesitius Moczar by its remarkably thick

antennae and easy to distinguish from the latter by lateral spines of propodeum

not stumpy, by head not converging behind eyes, by ocelli not in obtuse angle,

by size and proportions of some antennal joints, by tergite 2 with more scat-

tered punctures, etc.

REFERENCES

Moczar, L. 1970. Mesitinae of World, Genera Sulcomesitius Moczar and

Metrionotus Moézar Il. (Hymenoptera: Bethylidae). Acta Zool. Hung. 16:

40y-451.

Moczar, L. 1976. New Species of Sulcomesitius Moczar from Laos (Hymenop-

tera, Bethylidae). Ann. Hist. Nat. Mus. Natn. Hung. 68: 279-284.

P1.I-II.

Moczdr, L. 1977. A Review of the Genus Sulcomesitius Moczar (Hymenop-

tera: Bethylidae, Mesitinae). Acta Zool. Hung. 23: 139-170. PI.I-VIII.

ADDITION TO THE REVISION OF THE GENUS ATRACTODES

(HYMENOPTERA : ICHNEUMONIDAE) OF THE WESTERN

PALEARCTIC REGION I

Reijo Jussila

Krouvintie 7, SF-21330 Paattinen, Finland

Abstract

Atractodes thomsoni (Dalla Torre) is redescribed. New names are given

to two species whose earlier names are homonyms: A. thomsoni Jussila and

A. rufipes (Foerster). The former species gets the name A. townesi, n.

name and the latter A. fittoni, n. name. Atractodes tenuipes Thomson is

recorded from Britain for the first time.

This paper is the first addition to my revision of the genus Atractodes

Gravenhorst of the western Palearctic Region (Jussila 1979). The material for

this study was obtained from the collections of Department of Entomology,

University of Lund (EL), Zoologische Sammlung des bayerischen Staates, Munich

(XM) and R. Jussila (RJ).

1. Atractodes thomsoni (Dalla Torre) (Figs. 1-2)

Atractodes rufipes Thomson, 1884: 1025, F; holotype F: Sweden, Sk&ne,

Ringsjon (EL). A junior primary homonym of Atractodes rufipes

Provancher, 1884: 151 and Atractodes rufipes Foerster, 1876: 151, and

a junior secondary homonym of Atractodes rufipes (Foerster, 1876:

30) (Asyncrita).

Atractodes thomsonii Dalla Torre, 1902; 725. See Fitton 1982: 19.

Asyncrita thomsoni (Dalla Torre, 1902: 739) (Asyncrita). See Fitton

1982: 19.

This species is lacking in my revision of the western Palearctic Region

(Jussila 1979). It is redescribed below:

Female: Length 6.0 mm. Head shagreened on face and malar space, rest

of head polished, punctate on face; vertex not narrowed (but rounded) behind

compound eyes; temple broadest in its oral two-fifths; genal carina straight;

occipital carina rounded in its middle part; malar space about 1.0 x width of

mandible; clypeus rather convex, polished with punctured base, with middle

0.35 of its apical margin somewhat swollen, width about 3.0 x length; upper

tooth of mandible longer and wider than lower tooth; compound eye glabrous;

flagellum with 18 segments, length-to-thickness ratios: 2nd segment 3.5, 3rd

2.7, 5th 2.6, 7th and 10th about 1.8 and penultimate 1.1. Mesoscutum polished,

notaulus reaching about 0.3 the distance to scutellum. Mesopleuron polished,

somewhat shagreened beyond sternaulus; propodeum wrinkled, bearing rather

distinct carinae, median area parallel sided with transverse carinulae, length

about 2.3 x width, no apophysis or crest, propodeal spiracle small. Areolet

closed, second recurrent vein with two bullae; nervellus opposite. Legs not

very long and slender; length of hind coxa, seen from above, 1.8 x breadth;

length of hind femur 2.8 x width; claw think, about 1.5 x as long as arolium.

Abdomen long and laterally compressed from segment 2 to apex (Fig. 1); apex

of Ist segment, seen from side, straight (Fig. 2), cross section of petiole

depressed, length of postpetiole 1.5 x width; tergite | obscurely shagreened,

F = female. M = male. 501

202 Contrib. Amer. Ent. Inst., vol. 20, 1983

apically smooth, carinae weak, tergite 2 HOneHee length about 3.3 x width;

hypopygium without punctures.

Black; brownish yellow on mandible except for brownish teeth, and palpi

and tegula; yellowish orange on clypeus except for blackish base; yellowish

brown on stigma except for more transparent basal and apical corners. Legs

orange yellow.

Male : Unknown.

Material examined: The type-specimen of Atractodes rufipes Thomson.

An addition to the key to the females (Jussila 1979: 5):

23. Abdomen long and compressed from 2nd tergite on (Fig. 19 in Jussila

ar ae ea ees ee ee ee 23A

23A. Hind coxa long: length, seen from below, 2.5 x breadth; length of hind

femur 6.4 x width. Also flagellar segments long (e.g., Ist 5.0, 10th

2.4 and penultimate 2.0 x longer than broad). Clypeus, coxae, tro-

clanters, anc.micdic., and. hind. femoracblack . {025.60 < & 3 erie eres

ee ae a age lapponicus Jussila (p. 17 in Jussila 1979)

Hind coxa not long: length, seen from below, 1.8 x breadth; length of

hind femur 2.8 x width. Flagellar segments not long (e.g., Ist 3.5,

10th 1.8 and penultimate 1.1 x longer than broad). Clypeus partly

i Fe Se eC iy ei a’ Gece sb cee o thomsoni (Dalla Torre)

2. Atractodes townesi, n. name

Atractodes thomsoni Jussila, 1979: 14, F; holotype F: Sweden, Kjosen (EL).

The name thomsoni Jussila is a junior primary homonym of Atractodes

thomsoni (Dalla Torre). It is replaced by Atractodes townesi, n. name, to

honor Dr. Henry K. Townes.

3. Atractodes fittoni, n. name

Asyncrita rufipes Foerster, 1876: 14, F; lectotype F: Germany, Cologne,

by designation of R. Jussila, 1979 (ZM).

Atractodes rufipes, Jussila 1979: 31, F

The name rufipes (Foerster) is a junior secondary homonym of Atractodes

rufipes Provancher, 1874: 151 (Fitton 1982: 18-19). It is replaced by

Atractodes fittoni, to honor Dr. M. G. Fitton.

4. Atractodes tenuipes Thomson (Fig. 3)

Atractodes tenuipes Thomson 1884: 1022, M F; holotype F: Sweden,

Ostergotland (EL). The male specimens belong to Atractodes pauxillus

Foerster, 1876 and A. pusillus Foerster, 1876 (Jussila 1979: 20-21). Plesiotype

M: Sweden, Ang: Anger&n 10. VII.-20. IX. 1977 (K. Muller leg.) (RJ).

This species has earlier been tound only in Southern and Central Sweden.

But in August 1981 Mr. T. Bass collected some Limnophora riparia (Fallen

1824) (Dipt., Muscidae) pupae from a reservoir outflow, England, Dorset:

Wareham. This species of Atractodes emerged from some of the pupae. The

species is under these circumstances new in the British fauna. The family

Muscidae belongs to the cyclorrhaphous Diptera who are the only known hosts

of this genus (Jussila 1979: 3).

(holotype).

Figs. 1-5.

A. rufipes Thomson):

in sinistro-lateral view.

3, Atractodes tenuipes Thomson F (holotype). 4, A. pusillus Foerster, F

5, A. pusillus M (Sweden, Sk: P&lsjé).

203

1-2, Atractodes thomsoni (Dalla Torre), F (holotype of

1, Abdomen in dorsal view.

2, Petiolar segment

3-5, Apex of left hind leg in sinistro-lateral view:

204 ContribscAmer. Ent. fest... vel. 20, 1983

Among the British Atractodes species, A. tenuipes mostly resembles

A. pusillus Foerster, 1876 (1iogaster Thomson, 1884), but the hind tarsal

claw of the female shorter. Their apical part is as long as their basal part

(Fig. 3); in pusillus the apical part of the hind tarsal claws is shorter than

their basal part (Fig. 4). The male of tenuipes differs from pusillus male

by its claws, too, which resemble those of the female fet. Pig. 3). 208

claw of pusillus male, see Fig. 5. The abdomen of tenuipes is medially red-

dish but that of pusillus oftenest wholly dark.

ACKNOWLEDGMENTS

It is my pleasure to thank Mr. Hugo Andersson and Mr. Roy Danielsson,

Department of Entomology, University of Lund, for the loan of C. G. Thomson's

type specimens, and Dr. Erich Diller, Zoologische Sammlung des Bayerischen

Staates, Munich, for the loan of A. Foerster's Atractodes and asyncrita. col-

lection.

REFERENCES

Dalla Torre, C. G. de. 1902. Catalogus Hymenopterorum 3(2): 545-1141.

Lipsiae.

Fitton, M. G. 1982. A catalogue and reclassification of the Ichneumonidae

(Hymenoptera) described by C. G. Thomson. Bull. Brit. Mus. (Nat. Hist.)

45(1): 1-19.

Foerster, A. 1876. Synoptische ubersicht der Gattungen und Arten in der

Familie Stilpnoiden. Verh. Naturh. Ver. Rheinl. 33: 17-196.

Jussila, R. 1979. A revision of the genus Atractodes (Hymenoptera,

Ichneumonidae) of the western Palearctic Region. Acta. Ent. Fenn. 34:

1-44.

Provancher, L. 1874. Les Ichneumonides de Quebec avec description de

plusieurs especes nouvelles (part). Naturaliste Can. 6: 143-151.

Thomson, C. G. 1884. Forsok till gruppering och beskrifnig af crypti

(fortsattning), (Fasc. X). Opusc. Ent. 33: 939-1028.

: A NEW CLOUDFOREST TRACHYSPHYRUS FROM ECUADOR

AND COLOMBIA (HYMENOPTERA : ICHNEUMONIDAE)

Charles C. Porter *

Department of Biological Sciences

Fordham University, Bronx, New York 10458

Abstract

Trachysphyrus townesi, n. sp. may be recognized by its golden wings with

dark transverse bands, pale fulvous ground color, medially bituberculate clypeal

apex, narrow female postpetiole, and its strongly projecting ligulate female

propodeal cristae. It is one of the few Trachysphyrus to inhabit equatorial

cloudforests, the first species described from Colombia, and the third reported

from Ecuador.

INTRODUCTION

Trachysphyrus has about 200 species that are concentrated in Andean

and extratropical South America (Porter 1980: 29-32). Its major centers of

radiation include north Argentina (more than 100 species), Neantarctic Chile

(30-40 species), the Andes of Bolivia and Peru (25-30 species). One aberrant

Trachysphyrus has been cited from Guayana and Surinam. Only the

group, with a few species scattered throughout the Western Hemisphere from

Argentina to the northern United States, represents this genus outside South

America.

Trachysphyrus belongs to the Subtribe Ischnina (Townes 1969) of the

Tribe Mesostenini. It is closely related to the Holarctic genera Itamoplex

and Buathra, as well as to such basically Sonoran taxa as Compsocryptus,

Lanugo, and Joppidium. Townes (1969: 179-181) separated the Planosae group,

plus many other exclusively South American species, under the generic name

Chromocryptus Ashmead. Townes! viewpoint represents a conceptual advance

over the broader definition of Trachysphyrus offered in my 1967 revision of

the South American species. Trachysphyrus (sensu Porter 1967a) surely is

polyphyletic and susceptible of division into numerous genera. However, some

of its phyletic lines seem connected by annectant species. Consequently,

Townes' key (1969: 159-165) does not allow unequivocal routine identification.

Furthermore, during yearly fieldwork from 1968-1982 I have amassed more than

100 new species of Trachysphyrus from Argentina, north Chile, Bolivia, and

Peru. To attain a more rational generic classification of the "trachysphyroid"

complex, these undescribed species will have to be diagnosed and compared

with the 94 Trachysphyrus covered by me in 1967.

In the present contribution I describe a new and aberrant Trachysphyrus

from Colombia and Ecuador. Previously, only T. hoplites Porter and T.

metallicus Cameron had been known from Ecuador (Porter 1967b), and

Trachysphyrus never had been reported from Colombia. ‘The new species is

easy to recognize and phaenetically unique. However, its relationships to

other South American Trachysphyrus remain obscure. It may be a specialized

offshoot of the Rufitibia subgroup (Porter 1967a: 124), but it also shows

Research Associate , Florida State Collection of Arthropods at Gainesville.

205

206 Contrib. Amer. Ent. Inst., vol. 20, 1983

ESNet

‘

e ¢

Figs. 1-4. Trachysphyrus townesi, n. sp. 1, Fore

wing, female paratype. 2, Lateral view of ovipositor

tio; Female Bolotyoe, Sy Dorsal view of TiPrst terdi ze.

female paratype. 4, Lateral view of propodeum, female

Daratype.

Porter: Trachysphyrus (Ichneumonidae) 207

features reminiscent ot other "trachysphyroid" genera, particularly of Lanugo,

Compsocryptus, and Cryptopteryx.

I take special pleasure in dedicating this magnificent ichneumonid to

Dr. Henry K. Townes, Jr.

Trachysphyrus townesi, n. sp. (Figs. | -4)

Female: Color: Antenna pale fulvous on scape, pedicel, and flagellomeres

1-3, yellowish fulvous to yellow on tlagellomeres 4-15, and mostly blackish be-

yond flagellomere 15 or 16; head, body, and legs pale fulvous to yellowish ful-

vous with faint and irregular dusky staining that is best developed on front,

on vertex and upper temples or at least stemmaticum, on tegula interiorly,

on more or less of mesoscutum, on some thoracic sutures, sometimes on apex

of trochantelli and bases of femora, and on apices of some gastric tergites,

as well as with darker markings (black, deep brown, or sometimes light brown)

that are distributed as follows: on mandibular teeth; sometimes irregularly

and pallidly in dorsal trough of pronotum and on adjoining anterior meso-

scutum; narrowly and sometimes pallidly on anterior mesoscutum at base of

notaulus; sometimes briefly on mesoscutum above tegula; broadly near apex

of mesoscutum, in prescutellar depression, and on anterior scutellum; broadly

on prescutellar ridge; in most of meso and metanotal axillary troughs; briefly

on upper hind corner of pronotum; in much of area above subalarum; on an

extensive area that covers most of prepectus and which above expands rec-

tangularly onto anterior 0.2 of mesepisternum below subalarum; broadly on lower

hind corner of mesepisternum and more narrowly dorsad along hind margin of

mesepisternum to speculum; on all but dorsal 0.2 of mesepimeron; extensively

on mesosternum, with a median subtriangular projection reaching forward along

mesosternal sulcus; hind face of metanotum; throughout on upper metapleuron;

narrowly on front margin and more broadly on ventral margin and hind corner

of lower metapleuron; in trough between metanotum and propodeum and narrowly

along dorsal part of front margin of propodeum; broadly but irregularly or in-

completely on lateral propodeum between spiracle and base; on about apical

0.5 of crista; irregularly on most of lower 0.5 or more of hind face of pro-

podeum; on basal 0.8 of petiole above and laterally and almost throughout on

petiole below; on basal 0.4 of 2nd gastric tergite except for thyridium and

part of area between thyridium and base of tergite; on all but apical 0.2 of

3rd gastric tergite; more or less faintly on inner and outer dorso-apical

flanges of hind coxa; and on apical 0.5 of tarsal claws; fore wing with 3 dark

brown areas as follows: a broad dark cross band occupying most of apical

0.3 of median cell, basal 0.2 of discocubital cell, apical 0.3 of submedian cell,

basal 0.5 of Ist brachial cell, and adjoining part of anal cell; a similar brown

cross band covering basal 0.5 of radial cell, apical 0.3 of discocubital cell,

areolet, almost apical 0.5 of 2nd discoidal cell, and a little more pallidly ex-

tending across adjoining part of 2nd brachial cell; and with dark staining also

on tip of wing over apical 0.5 of 3rd cubital cell, broadly above and more

narrowly ventrad on apex of 3rd discoidal cell, and narrowly on apical margin

of wing below 3rd discoidal cell; hind wing with a dusky cloud toward apex

of brachiellan cell and with apical 0.2 more conspicuously blackish; wings

otherwise golden yellow.

Length of fore wing 12.0-12.5 mm. Flagellum filiform, very long and

slender, not flattened below toward apex; Ist segment 5.6 -6.4 as long as deep

208 Contrib Amer Ent. tnsty;: vol: 20; 1983

at apex. Clypeus in profile strongly convex and a little asymmetric, with high-

est point distad of middle and with apical face steeper than basal; apical mar-

gin thin and weakly reflexed, especially laterad, and with the apex itself

broad, slightly convex, and interrupted on each side of the middle by a weak

tubercle. Malar space 0.95 -0.97 as long as basal width of mandible. Temple

strongly and directly receding in dorsal view; in lareral view 0.36 -0.38 as

long as eye. Occipital and hypostomal carinae: Occipital carina sharp and

weakly elevated throughout, joining the only slightly more elevated hypo -

stomal carina well above base of mandible. Pronotum: Dorsal submarginal

groove obsolete; dorsal margin not swollen; epomia absent above scrobe, strong

in scrobe, and prolonged some distance below onto low front swelling of pro-

notum. Mesoscutum convex; notaulus very faintly traceable for 0.5 -0.6 the

length of mesoscutum, not impressed; surface mat with abundant, superficial,

minute punctures that emit close-packed, short, recumbant setae. Meso-~

pleuron: Subalarum flattened and weakly ledge-like, crescentic in exterior con-

tour, subcarinate on outer margin; surface mat and minutely granular with

speculum a little swollen but similar in sculpture to rest of mesopleuron; pre-

pectus without a ridge opposite lower hind corner of pronotum; sternaulus

strong basally but becoming abruptly obsolete about 0.5 the distance between

prepectal carina and hind corner of mesopleuron. Wing venation: Radial cell

3.9-4.6 as long as wide; areolet large, symmetrically pentagonal, intercubiti

strongly convergent above, 2nd abscissa of radius 0.7 as long as Ist intercubi-

tus; discocubitus gently and evenly arched, without a ramellus; 2nd recurrent

broadly angled at bulla, moderately inclivous above bulla and almost vertical

below it; nervulus about 0.20 its length antefurcal, gently inclivous; mediella

almost straight on apical 0.6, arched basad; nervellus broken far below middle,

upper segment 2.0 -3.0 as long as lower; axillus paralleling hind margin of wing

at about 0.4 the distance between hind margin and submediella, near apex be-

coming weaker and sometimes curved toward hind margin. Fore tibia elongate,

faintly inflated. Hind coxa with a strong, polished subvertical groove located

externoventrally near base. Propodeum comparatively short and high; apical

face sharply discrete from basal and almost vertical, 1.0 as long as basal;

spiracle elongate, 3.6~4.0 as long as wide; basal trans-carina on dorsum trace-

able but weak to partially effaced, broadly curved forward medially, becoming

strong laterad and sublaterally raised into a small, crescentic crista; apical

trans—carina represented only by its unusually large and strongly projecting,

narrowly ligulate cristae, which are 0.5 or more as long as apical face of pro-

podeum; longitudinal carinae absent, except for a foveolate trace of pleural

carina; surface mat with tiny, exceedingly dense punctures emitting innumer-

able, very short, crowded setae that are recumbant on dorsum and tend to be

more erect elsewhere, as well as with weak wrinkling along basal trans-

carina, variably developed faint transverse wrinkling apicad on basal face and

with faint to well developed transversely to longitudinally biased wrinkling on

apical face, that is fine to obsolete above and which becomes at least a lit-

tle stronger ventrad. Ist gastric tergite: Petiole at base with a strong,

triangular lateral tooth; postpetiole rather elongate, almost parallel-sided, mod-

erately expanded, 0.88 -1.0 as wide apically as long from spiracle to apex;

ventro-lateral carina absent, except on postpetiole; dorso-lateral carina faintly

traceable throughout; dorsal carinae weakly suggested above spiracles on apex

ot petiole and base of postpetiole; surface of postpetiole mat with fine,

minutely granular puncto-reticulation and very dense, short, recumbant setae.

Porter: Trachysphyrus (Ichneumonidae) 209

2nd gastric tergite uniformly mat, similar in sculpture and setation to post-

petiole; thyridium large, subcircular, removed by a little more than its own

length from base of tergite. Gaster elongately fusiform; succeeding tergites

similar to 2nd but reareward with progressively finer micro-sculpture and in

part dully shining. Ovipositor : Sheathed portion 0.48-0.55 as long as fore.

wing; straight, moderately stout, compressed; nodus faintly suggested, its po-

sition marked by a tiny notch; dorsal valve with a long and gently convex

taper between notch and apex, taper accentuated on apical 0.3 of tip; ventral

valve on tip with fine, well space, inclivously oblique ridges; tip 0.18 -0.21 as

high at notch as long from notch to apex.

Male: Differs from female as follows: Color: Scape and pedicel pale

fulvous; flagellum pale fulvous on segments 1-10 or 11, becoming tinged with

yellow on segments I] or 12-17 or 18, yellowish fulvous grading into black on

17 or 18, and black on succeeding segments; with black variably developed in

mesonotal axillary trough; metanotal axillary trough dull fulvous with brown

Staining; black area below subalarum sometimes more irregularly shaped than

in female, sometimes margined with brownish; dorsal 0.3 of mesepimeron pale

fulvous; upper metapleuron with a large, ovoid fulvous blotch; lower meta-

pleuron without any black along ventral margin; spiracular area of propodeum

more restrictedly black than in female; no black on propodeal cristae; hind

face of propodeum sometimes largely black with a median extention onto dor-

sal face; Ist gastric segment mostly fulvous with blackish irregularly on basal

0.3 or less of petiole above and dorso-laterally and on about basal 0.5 ventro-

laterally and ventrally; sometimes with blackish on basal 0.3 of 3rd tergite

Or sometimes with tergites 2 and following almost uniformly fulvous and with-

out well defined darker markings, although with some irregular and faint

dusky staining; dark bands of fore wing sometimes a little paler than in fe-

male, with the apical blotch broader and the 2 cross bands a little narrower.

Length of fore wing l1.5 -l1.9 mm. Flagellum weakly tapered toward apex

and slightly stouter than in female; segments with short, dense, uninterrupted

erect setae below; tyloids developed as fine longitudinal ridges on flagellomeres

18-21, those on 18 and 19 nearly percurrent, that on 20 shorter, and tyloid on

21 very short; Ist segment 3.5 as long as deep at apex. Clypeus in profile less

strongly raised and more asymmetric than in female, basal face gently con-

vex, apical face short and steep, highest point of clypeus closer to apex than

in female. Malar space 0.72 -0.80 as long as basal width of mandible. Tem-

ple: 0.43 -0.48 as long as eye in lateral view. Pronotum: Epomia present

only in scrobe, sometimes weak. Mesoscutum similar to female but with the

extremely dense setae longer and inconspicuously erect. Mesopleuron: Outer

margin of subalarum more sharply carinate than in female; surface dully

shining with fine and dense punctures and, on area between prepectus and sub-

alarum, with extensive fine wrinkling that is weak and irregular below but be-

comes more or less stronger and somewhat longitudinally biased dorsad and

dorso-anteriad; speculum finely punctate but not wrinkled. Propodeum in pro-

file gently, convexly, and almost evenly sloping between base and apex; basal

and apical faces not sharply discrete; apical face 0.67 -0.73 as long as basal;

spiracle 2.8 -3.0 as long as wide; apical carina represented only by small,

triangular, moderately projecting cristae; surface dull to, in places, bright~

ly shining, with much fine wrinkling and abundant small, dense punctures,

which are more discrete than in female, and with conspicuous, very dense,

210 Contrib. Amer. Ent. Inst., vol. 20, 1983

moderately long, erect setae; pleural carina more or less completely defined

between spiracle and apex. Ist gastric tergite: Petiole at base with a weak,

subtriangular lateral expansion; postpetiole weakly expanded, elongate, 0.61 -

0.66 as wide apically as long from spiracle to apex; dorsal and dorso-lateral

carinae scarcely traceable; surface of postpetiole shining with abundant, mod-

erately small, shallow punctures that become sparser on apical 0.5 and which

emit long, semirecumbant, extensively overlapping setae. 2nd gastric tergite

dully shining with tiny, dense but discrete punctures that emit rather long,

semirecumbant, much overlapping setae. Gaster elongate and slightly fusi-

form; succeeding tergites similar to 2nd but less shining and with even denser

minute punctures and strongly overlapping setae.

Holotype: Female, Ecuador, near Baeza at about km 10 on road to Tena,

1800 m, June 8, 1979, C. Porter (CPH, Gainesville). Paratypes: Female,

Ecuador, same data as type (Gainesville); 2 males, Colombia, Valle, km 8

camino Cali-Dagua, 1800 m, Aug. 29, 1975, L. Stange (Gainesville, Porter).

The holotype will be deposited in the collection of Henry K. Townes, now

being transferred to the Center for Parasitic Hymenoptera at the Department

of Entomology and Nematology, Unverisity of Florida, Gainesville, Florida.

One female and | male paratype will be placed in the Florida State Collection

of Arthropods in Gainesville at the Division of Plant Industry of the Florida

Department of Agriculture and Consumer Services. The 2nd male paratype

is retained for the author's personal collection (301 North 39th Street, McAllen,

Texas 78501).

Relationships: Trachysphyrus townesi fits the definition of Trachy-

sphyrus as provided by Porter (1967a: 37-38). Its dordally convergent inter-

cubiti, angled 2nd recurrent, and slender habitus might relate T. townesi to

the Rufitibia subgroup of Trachysphyrus (Porter 1967a: 124-125). Indeed,

T. hoplites Porter (1967b) of this subgroup also occurs in Ecuador and shares

with T. townesi such specialized features as its gently reflexed clypeal mar-

gin, crescentically lamellate subalarum, and unusually long propodeal cristae.

T. townesi differs from all other Trachysphyrus by its golden wings with

dark cross bands, fulvous ground color, and by the following combined struc-

tural features: clypeus with a pair of tiny submedian tubercles on apical mar-

gin; notauli extremely faint; areolet with intercubiti strongly convergent above;

2nd recurrent broadly angled at bulla; mediella almost straight on apical 0.6;

propodeum in female high with basal and apical faces subequal and sharply

discrete but in male gently and evenly sloping between base and apex, with

basal trans-carina weak, longitudinal carinae almost absent, and apical trans-

carina absent except for its very large and strongly projecting ligulate cristae

in the female and small, triangular cristae in the male; propodeal spiracle

2.8-4.0 as long as wide; base of female petiole with a large, triangular later-

al expansion, of male petiole with a weak and ledge-like excrescence; post-

petiole long and little expanded rearward, in female 0.88-1.0 as wide at apex

as long from spiracle to apex, in male 0.61-0.66 as wide; 2nd gastric tergite,

and most of body, mat with dense, minute punctures and very dense, tiny setae

that are mostly recumbant in female and partly erect in male; ovipositor

straight, rather robust, compressed, with nodus indicated only by a minute notch

and apex, tip 0.18 -0.21 as high at notch as long from notch to apex.

Porter: Trachysphyrus (Ichneumonidae) 211

Because of its dark-banded wings and fulvous ground color, T. townesi

superficially resembles many species of Lanugo and Compsocryptus. These 2

close relatives of Trachysphyrus are centered in the Sonoran Region, but have

also a few species from Ecuador and elsewhere in South America.

All Compsocryptus differ from T. townesi in having the 2nd abscissa of

the radius as long as the Ist intercubitus; the axillus distant from the hind

margin of the wing; the propodeal cristae of both sexes relatively low; the

ventro-lateral carina strong and complete on the Ist gastric segment; and the

Oovipositor upcurved.

Species of Lanugo may be separated from T. townesi_ by their lack of

tubercles on the clypeal apex; weakly grooved hind coxal base; propodeum with

at most moderately large cristae and with the apical trans-carina more or less

well defined throughout; and female postpetiole that is usually much wider at

apex than long from spiracle to apex. In many features, however, T.

townesi seems close to Lanugo. It might be related to the stock from which

that genus was derived.

Cryptopteryx, a cloudforest "trachysphyroid" genus of Andean South

America superficially resembles T. townesi in its very large and projecting

propodeal cristae and mat, densely micro~punctate and micro-setose body sur-

face. As mentioned by Townes (1969: 180), cryptopteryx easily could be sy-

nonymized with Trachysphyrus. However, it is not directly related to T.

townesi. Species of Cryptopteryx have no clypeal tubercles and the clypeus

nasutely elevated in profile; have the pronotal dorsum shallowly grooved; the

notauli moderately impressed; the epomia prolonged and curved mesad above

scrobe; the intercubiti weakly convergent above; the discocubitus broadly

angled with a prominent ramellus; the axillus very close to the hind margin

of the wing; the propodeal cristae apically sharp, thorn-like, and equally de-

veloped in both sexes; the propodeum more completely areolated with its longi-

tudinal carinae extensively traceable; the Ist gastric segment with a weak

baso-lateral flange in both sexes; and the ovipositor tip sagittate.

Field notes: The holotype and paratype females were netted from

undergrowth in east Andean cloudforest about 100 km from Quito, Ecuador.

Here the overstory trees (Myrtaceae, Lauraceae, etc.) support a lush

epiphyton of orchids and bromeliads, tree ferns grow abundantly, and the

forest floor is covered by a dense herbaceous stratum. This kind of habitat

is optimum for parasitic Hymenoptera and yielded more than 100 ichneumonids

in less than 2 hours. Among the "trachysphyroids" obtained at this locality

in addition to T. townesi were 3 females and 3 males of Cryptopteryx

columbianus. C. columbianus previously has been reported only from Colombia

(Townes 1966: 75).

ACKNOWLEDGMENTS

My 1979 fieldwork in Ecuador was supported by a grant from the Com-

mittee for Research and Exploration of the National Geographic Society. The

Division of Plant Industry of the Florida Department of Agriculture and Con-

sumer Services generously furthered this and related projects by provision of

collecting supplies, xeroxing of rare publications, and in many other ways.

212 Contrib. Amer. Ent. Inst., vol. 20, 1983

Among the staff of that institution, Mr. Harold Denmark, Dr. Howard V.

Weems, Jr., and Dr. Lionel A. Stange were particularly helpful.

REFERENCES

Porter, C. 1967a. A revision of the South American species of

Trachysphyrus. Mem. Amer. Ent. Inst. 10: 1-367.

Porter, C. 1967b. A new species of Trachysphyrus from the Andes of Ecuador.

Psyche 74: 27-33.

Porter, C. 1980. Zoogeografia de las Ichneumonidae latino-americanas. Acta

Zool. Lilloana 36: 5-52.

Townes, H. K. 1966. A catalog and reclassification of the Neotropic Ichneu-

monidae. Mem. Amer. Ent. Inst. 8: 1-367.

Townes, H. K. 1969. Genera of Ichneumonidae, Part 2: Gelinae. Mem. Amer.

Ent. Inst. 12> 1-537.

A NEW SPECIES OF THE GENUS KARPINSKIELLA

(HYMENOPTERA : CHALCIDOIDEA : PTEROMALIDAE)

Karl-Johan Hedgqvist

Swedish Museum of Natural History, Entomology Section

S-104 05 Stockholm, Sweden

Karpinskiella townesi, n. sp. is described from Brazil. A key

to the known species of the genus is given.

The genus Karpinskiella was described by Boucek in 1955 with one

species K. pityophthori Boué. In the year 1968 Hagen and Caltagirone

described a second species K. paratomicobia Hag. and Caltag. A new species,

K. townesi, n. sp. is described here from Brazil. The latter extends the

range for the genus from the Holarctic Region into the Neotropical Region.

Probably the genus has a worldwide distribution as parasitoids on the bark

beetles.

Karpinskiella townesi, n. sp.

Female: Dark blue with dark coppery reflection on vertex and dorsally

on thorax; propodeum with tint of green and gaster with brownish reflection.

Scape, pedicellus and I:st annellus yellowish brown. The rest of antenna dark

brown. Legs yellowish brown with mid and hind coxae partly brownish, mid

femora with more and less brown reflection, hind femora dark brown and all

clawjoints brown. Tegulae brown. Wing veins pale yellowish brown.

Head (Fig. 2C) transverse, distinctly wider than thorax. Clypeus dis-

tinct incised in the middle. Antennae, 11353, (Fig. 3B) inserted above a level

with anterior margin of eyes. Antennal scape reaches above median ocellus.

Pronotum with collar in the middle with a carina. Head and thorax

reticulated with medium large meshes on head and behind frenum of scutel-

lum. Prepectus of medium size. Propodeum with a medium carina. Gaster

short, shorter than thorax. Legs strong with all femora swollen. Posterior

margin of l:st tergite incised in the middle.

Fore wing (Fig. 3F) with long marginal vein, three times longer than the

stigmal vein.

Male: Unknown.

Length: °253 mm * 25" mm.

Holotype: F, Brazil, Nova Teutonia (27° ll' B 55° 23' L), 27.VIL.1962,

leg. Fritz Plaumann. (Coll. Hedqvist). Paratype: 3 F from the same lo-

cality, 5.VIII.1945, leg. Fritz Plaumann. (Coll. Hedqvist).

The description of the genus Karpinskiella Bout. is modified to include

K. paratomicobia Hag. and Caltag. and K. townesi, n. sp.

Genus KARPINSKIELLA Boucek

Head stout, subglobose to transverse, distinctly broader than thorax.

Facial cavity above antennal sockets small but distinct. Antennae, 11353, in-

serted above the lower ocular line (Fig. 2A-C). Flagellar joints transverse to

longer than wide, scape not reach or reach median ocellus. Clypeus

213

214 Contrib. Amer. Ent. Imst., vol. 20, 1983

moderately but distinctly incised in the middle of anterior margin. Mandibles

with 3 + 3 teeth. Malar space large.

Pronotum with collar more and less separated. Notauli incomplete.

Scutellum longer than wide with frenum indistinct. Propodeum with or with-

out median carina and more and less distinct plicae. Prepectus from small

to medium size. Gaster long-oval as long as or shorter than thorax, a little

compressed from the sides (F). Legs with femora more and less swollen,

hind tibia with one spur.

Wings naked in basal third, the rest covered with sparse short hairs,

ciliated on the margin. Marginal vein one and half times longer to three

times longer than stigmal vein, postmarginal vein a little longer than stigmal

vein.

Type-species: Karpinskiella pityophthori Bouc, by monotypy.

KEY TO SPECIES OF THE GENUS KARPINSKIELLA

1. Antennal scape short not reaching to median ocellus, funicular segments

transverse. Collar indistinct. Propodeum without median carina (Figs.

L. 28, 2A. and D) Finland, Norway, .Polen,..Sweden. .......... ... =...

Beaea tien gt sy Mus einai wR | Sse ee Ca REG le Ue ge i Ke DLEBOORTHOrI

Antennal scape longer and funicular segments longer than broad. Collar

Gistinei 4 ropodenm, «withia \Median,.CaLina. & oie i. lene. Sage cee cme = ae

2. Scutellum flat. Collar rounded. Marginal vein one and half times longer

than stigmal vein (Figs. 2A, 3C and E) Utah, Arizona, California .. .

SE eee ee ee Men a 6 LINO, te SNE gc Cg Mn nee aeRO RReRERrE OU cee K. paratomicobia

Scutellum more vaulted. Collar anterior with a carina. Marginal vein

three times as long as stigmal vein (Figs. 2C, 3B and F) Brazil ....

. K. townesi

REFERENCES

Bouéek, Z. 1955. Parasiti kuoroce Pityophthorus polonicus Karp. z pieniu.

Roczn. Nauk. les. Il: 83-92.

Hagen, K. S. and Caltagirone, L. E. 1968. A new species of Karpinskiella

(Hymenoptera: Pteromalidae). Pan-Pacific Ent. 44: 241-248.

Hedgqvist, K.-J. 1963. Die Feinde der Borkenkafer in Schweden. I Erzwespen

(Chalcidoidea). Studia Forestalia Suecica. No. ll, 176 pp. Stockholm.

Hedaqvist: Karpinskiella (Pteromalidae) 215

le om >

)

Fig. 1, Karpinskiella pityophthori Boucek, female.

216 Contrib. Amer. Ent. Inst., vol. 20, 1983

Fig. 2. A, Karpinskiella paratomicobia Hag. and Caltag. F, head in

front view. B. K. pityophthori Bou. F, head in frontal view. C, K.

townesi, n. sp. F, head in frontal view.

Hedqvist: Karpinskiella (Pteromalidae) aii

Fig. 3. A, Karpinskiella pityophthori Bouc. F, antenna. B, K. townesi,

n. sp. F, antenna. C, K. paratomicobia Hag. and Caltag. F, antenna. D, K.

pityophthori F, marginal, stigmal and postmarginal vein of fore wing. F, K.

townesi, n. sp. F, marginal, stigmal and postmarginal vein of fore wing.

STUDIES ON HYMENOPTERA ICHNEUMONIDAE

IX. ‘THE RUDOLF JUSSEL'S COLLECTION

Franco Frilli

Istituto di Difesa delle Piante

Universita degli Studi - Udine, Italy

In the latest decades many researchers have been studying the old

Ichneumonidae collections in order to get and study types, which are neces-

sary for revisions of genera or various systematic groups. Moreover, some

colleagues are very interested in Jussel's Ichneumonidae collection which is

in Italy at present.

We know very little about Rudolf Jussel, the entomologist who taught

at Bregenz (Austria). His only publication on Ichneumonidae "Beitrage zum

voralberg'schen (ésterreichischen) Insektenfauna" show that in 1907 (the date

of its issue) he owned an entomological collection of 3700 species, as the re-

sult of his 16 years collecting.

Horn and Kahle (1935-37) mentioned that in 1932 the sawflies of that

collection reached A. Dodero in Genoa, through A. Winkler. The Diptera were

given to "Pietro Rossi" Entomological Museum of Duino (Trieste)* The remain-

ing Hymenoptera specimens were included in Albert Winkler's private collection

in Vienna in about 1925. |

In 1945, when Albert Winkler died, his Hymenoptera collection was be-

queathed; later on, his heirs sold it. In 1960 Universita Cattolica del Sacro

Cuore (Italy) bought all the available Ichneumonidae of Winkler's collection

for the Entomological Institute of Agriculture Faculty at Piacenza: and they

are still there. Most of the identified specimens belonged to R. Jussel's col-

lection. They are easily identifiable because they are prepared and labelled

in an uniform way. Beginning from the top, each specimen has three labels:

the first one points out the sex and date of collecting;

the second one points out the place of collecting;

the third one has a black frame and points out the identification.

The specimens coming from other collections (it is easy to notice that

many of them were previously included in O. Schmiedeknecht's, O. Staudinger's

and R. Schmidt's collections) have the black framed labels with the identifica-

tion written by Jussel, but the other indications are missing or are different.

Sometimes we can find also labels with the identification by E. Clement, O.

Schmiedeknecht and others.

We cannot be sure whether the collection of Piacenza includes all the

Ichneumonidae of Jussel's original collection or if there were intentional or

accidental reductions. Some of Jussel's Ichneumonidae were probably lost or

are elsewhere. Some of the specimens mentioned in Jussel's work (1907)

(e.g., Ichneumon patruelis Holm., Lissonota antennalis Thoms., etc) are

not there, and among them some specimens which were certainly included in

Jussel's collection (see Schmiedeknecht, 1902-36, I, p. 212 for Pyramidophorus

flavoguttatus Tisch. and Ill, p. 1208 for Glypta lineata Desv.).

—— ee ne ee es

* Poday in Mugeo Civico di Storia, Naturale, Minano CPE Big}.

218

Frilli: Ichneumonidae 219

Ichneumoninae: 388 specimens belonging to 172 species;

Cryptinae: 256 specimens belonging to 138 species;

Pimplinae: 222 specimens belonging to 86 species;

Ophioninae: 359 specimens belonging to 192 species;

Tryphoninae: 316 specimens belonging to 183 species.

The specimens are very well kept, mostly collected in Austria; but there

are also specimens from Syria, Tunisia, etc.

It is certain that other specimens of different origin were added to the

ones labelled by Jussel. Some specimens have indications similar to Jussel's.

Many others were probably added by Winkler. There are 217 specimens that

were partly added to already existing species and partly belong to 62 species

which Jussel did not have (there are no specimens with Jussel's labels). These

are:

Added specimens Species probably not owned

by R. Jussel

Ichneumoninae 64 23

Cryptinae 16 5

Pimplinae 98 2%

Ophioninae 37 12

Tryphoninae ee re

217 62

The following species are of some significance:

"Euthycryptus scrobiculifer n.g., n. sp."

Jussel, 1907. Voralb. Landes-Mus. Bregenz, Jahresber. 44: 71. F.

It is the only specimen which perfectly correspond to the description.

It is certainly the holotype. "23.7, F", "Hochgerach", "Eutycryptus scrobi-

culifer Jussel". Horstmann (in litteris) is planning a revision of

genus group, which will facilitate its identification.

"Ghathophya mediterranea Schmiedeknecht"

Schmiedeknecht, 1900. Term. Fuzetek 23: 239. M, F.

There are four specimens (1 F and 3 M) with only one identification

label by Schmiedeknecht: "Ghathophya mediterranea F, M, Schmiedeknecht

Tunis". Therefore, the collecting place is Tunisia.

Schmiedeknecht quotes south Italy, Algeria and Tunisia as "Terra typica".

The species (according to Oehlke and Horstmann, in litteris) is also present

in Berlin Museum (1 M and 1 F); but no designation of types has been pub-

lished till now. Correct name: Odinophora mediterranea (Schmiedeknecht).

"Hemiteles n. sp."

One specimen with Jussel label "Hemiteles n. sp. Jussel". Correct

name: Hemiteles vicinus (Gravenhorst) (Teste Horstmann, in litteris).

220 Contrib. Amer. Ent. Inst., vol. 20, 1983

REFERENCES

Aubert, J. F. 1978. Les Ichneumonides ouest-palearctiques et leurs hotes.

OPIDA, E-chauffour.

Horn, W. and Kahle, J. 1935-37. Uber entomologische Sammlungen, Ento-

mologen und Entomo-museologie. Ent. Beihefte aus Berlin-Dahlem 2-4:

1-536.

Jussel, R. 1907. Beitrage zur voralberg'schen (osterreichischen) Insekten-

fauna. "Festschrift zum 50jahrigen Bestande zugleich 44. Jahresbericht

uber das Jahr 1906", Bregenz: 67-73.

Schmiedeknecht, O. 1900. Neue Hymenopteren aus Nord-Afrika. Term.

Fuzetek. 23: 234-247.

Schmiedeknecht, O. 1902-1936. Opuscula Icneumonologica. Blankenburg. 5

voll. + 25 suppl.

Townes, H. and Townes, M. 1962. Ichneumon-flies of America North of

Mexico. 3. Subfamily Gelinae, Tribe Mesostenini. U.S. Natl. Mus. Bull.

216, 602 pp.

DESCRIPTIONS OF SOME NEW AFRICAN SPECIES

OF OPIUS FROM THE TOWNES COLLECTION

(HYMENOPTERA : BRACONIDAE : OPIINAE)

Maximilian Fischer

Natural History Museum, Vienna, Austria

Abstract

Keys are proposed for identification of 14 species of the subgenus

Grimnirus Fischer and 11 Old World species of the subgenus Frekius Fischer

(genus Opius Wesmael, Hym., Braconidae, Opiinae). The following species

are described as new: Opius (Grimnirus) eutownesi, ileifensis, nigeriensis,

obuduensis, toromojaensis, and Opius (Frekius) nairobicus. Important

morphological details are figured.

Among the Opius in the Townes Collection there are some new species

belonging to the subgenera Grimnirus Fischer and Frekius Fischer. Below

they are described in full. I make use of the occasion to revise the Old World

species of the subgenera mentioned, and propose new keys for identification

of the species involved.

The following abbreviations are used: AEI = American Entomological

Institute; rl, r2, r3 = abscissae of radial vein (r); ccl, cc2 = cubital cross

veins; d = discoidal vein; nr = recurrent vein; np = parallel vein, R = radial

cell; Cu2 = second cubital cell; B = brachial cell.

I wish to thank Dr. Henry K. Townes very much for all the valuable as-

sistance he granted me in my research work on the ape MmaNes of the

Braconidae.

Subgenus GRIMNIRUS Fischer

Subgenus Grimnirus Fischer, 1971. Ann. Naturh. Mus. Wien 75: 389, 39].

Type-species: Opius fuscicarpus Szépligeti (designated by Fischer

1972). Appeared actually 1972. Fischer, 1972. Das Tierreich 91: 70

Fischer, 1973. Polskie Pismo Ent. 43: 242 (key to Ethiopian species).

Mesoscutum with dorsal fovea, precoxal suture sculptured, mouth closed,

mandibles not expanded at base, clypeus without tooth on lower edge, head

dorsally smooth or at most occiput punctate, metasoma at least with fine

sculpture on tergite (2 + 3), nr postfurcal.

All species of this subgenus are known from Ethiopian Africa except

O. (G.) durigaster Fischer which is from Bismarck Islands.

KEY TO THE SPECIES OF OPIUS (GRIMNIRUS)

1. Thorax 413 simeeiae dane as ies Ue a ee ere 2 ri

Whorax’ ativmose 43) timed as dong de high 6). eee Lee ae es: 5

2. Sternaulus crenulate and rugose. Sheaths of terebra of female rather

thin. Legs and body predominantly dark ... 4. obuduensis, n. sp.

Sternaulus weakly impressed, only shagreened. Sheaths of terebra of

female rather broad. Legs entirely yellow, body with yellow pat-

tern

222

10.

11.

12.

Lo.

Contrib. Amer. Ent. Inst., vol. 20, 1983

Thorax 1.4 times as long as high. Face punctate and shagreened. 4.0

mim. Miri 2ealre oa er ek congoensis Fischer

Thorax 1.5 times as long as high. Face evenly, fine shagreened.... 4

Length of body 4.3 mm. Head, thorax and base of metasoma dark.

Metasomal tergite 1 a little shorter than apically wide. F.

WIENS (ea oe ee ee ee fuscicarpus Szepligeti

Length of body 2.8 mm. Body entirely bright yellow. Metasomal ter-

site lias lone as apically wide. Fi. Botswana: -.: sc. kate. «

Gu be eee Ge ee ha We ee ee oa 2 5... Eoromojaensis, mn.* sp.

MEsipleuiom SMOGLN . 4.010). Bo oe. ie Pe ee 6

Mesoplevron: sharteenédon disease) ork) eke. Yin es seen 10

Metasomal tergite 2 regularly and densely longitudinally striate. 2.0 mm.

fo Bismarck telands:. 2 cae ke ee es durigaster Fischer

Metasomal tergite 2 shagreened to feebly rugose ............ 7

Clypeus at least twice as wide as high, lower edge straight. 2 mm.

Mi. NIg@TiA ae ie a ae ea l.. eutownesi, fn. sp.

Clypeus about 1.7 times as wide as high, lower edge convex ...... 8

Head 1.9 times as wide as long, eyes 1.8 times as long as temples. 2.4

Mit NP aie ea ee a a a es leroyi Fischer

Head about 2.2 times as wide as long, eyes 2.5 times as long as

PO ee ee ee a ae Be eee ee ea a meg ae 7

Metasomal tergite 1 1.3 times as long as apically wide. Hind femora 5

times as fong as wide. 1.6 mm. Burundi, M, F. Zaire... -.. . +

ea ke ee eC ek ee are era Gye meta oa at Gy bururianus Fischer

Metasomal tergite 1 as long as apically wide. Hind femora 4 times as

fong as wide. 18 mm. F,,..Nigetia ... ..,3.. nigeriensis, nN. Sp.

Head and metasoma bright yellow, the latter especially on basal half.

3.0, Mie OF eouthOACa yy. 6 ed palligaster Fischer

Out ONC ely DCR a i a a ee Ge Gy ee a ea 11

Mesoscutum delicately shagreened. 2.4 mm. M. Uganda........

eRe ea te a are, er mujenjensis Fischer

Meso sCUTUM CNTircly SINOOLN Ge oe oo a ek ee Shae ae bee waa ee 12

Metasomal tergite 1 longitudinally striate, tergites 2 and 3 evenly,

delicately granulate. Antennae as long as body, 20-jointed. [Head

2.4 times as wide as long.) 2.0 mame «Ff. bouth Aftiog ys 26 arias:

Nay SRN ee re EORTC SO cr PO GCL iG amen minicornis Fischer

Metasomal tergite 1, 2 and part of 3 evenly, delicately coriaceous to

rugose. Antennae 1.5 times as long as body, 25- 27-jointed. [Head

Pe 2 ies, a Wee ne hi aa al, uk ete oe ees 13

Ocelli arranged in an isosceles triangle, the base of which is longer than

a side. Cu2 rather parallel sided, ccl 1.8 times as long as cc2.

Metasomal tergite 1 as long as apically wide, delicately and evenly

cOnlaceous-rugose. 1.6 mm. M.° Uganda, South Africa . 4)... -

Be We tase UR ag Re gl aah Soda caer aay eto ae Oe ake katonicus Fischer

Ocelli arranged in an equilateral triangle. Cu2 distad conspicuously

narrowed, ccl 2.2 times as long as cc2. Metasomal tergite 1 1.25

M = male. F = female.

Fischer: Opius (Braconidae) 223

times as long as.apically wide, shining. 1.9 mm. F... Nigeria . ...

Ug a heer ees ne Sie Mun seed coe cael ee 2. ileifensis, Nn. sp.

1. Opius (Grimnirus) eutownesi, n. sp. (Figs. | - 3)

Because of the rather short thorax, the shining mesopleura, the hidden

terebra and the lack of striae on metasomal tergite 2, close to Opius leroyi

Fischer, O. bururianus Fischer and 0. nigeriensis, n. sp. Opius eutownesi,

n. sp. is distinguished from them by the comparatively broad clypeus (2.1 times

as wide as high) and the straight lower edge.

Female: Head, the major part of the thorax and the postrior half of the

metasoma shining. Head 2.2 times as wide as the face, 2.1 times as wide as

metasomal tergite | posteriorly; only occiput with inconspicuous hairs; eyes

protruding, eyes and temples commonly rounded, eyes 2.4 times as long as the

temples, the distance of the toruli from each other greater than the distance

from the eyes, occiput sinuate; ocelli strongly protruding, the distance between

them little more than the diameter of an ocellus, distance between an ocellus

and the edge of an eye as long as the width of ocellar area. Face 1.1 times

as wide as high, sparsely puntuated and with minute hairs, median polished

area weakly developed, margins of eyes nearly parallel sided. Clypeus 2.1 times

as wide as high, flat, separated from the face by an evenly curved suture,

lower edge straight. Distance of tentorial pits from each other 2.5 times as

long as distance from the eyes. Genae shorter than the width of the mandibles

basally. Mouth closed, mandibles not expanded basally, maxillary palpi a trifle

longer than the height of the head. Eye in lateral view 1.8 times as high as

long, 1.5 times as long as the width of the temples, temple parallel sided.

Antennae 1.4 times as long as body, 27-jointed; flagellar joint | three times,

joint 15 and the penultimate twice as long as wide; hairs shorter than width

of flagellar segments, in lateral view 3 -4 sensillae visible. Thorax 1.33 times

as long as high, 1.3 times as high as the head, upper side strongly convex.

Mesoscutum 1.25 times as wide as long. In front of the tegulae rather evenly

rounded, middle lobe only a little protruding; notauli anteriorly deep, weakly

crenulate, reaching on the disc, there disappearing, dorsal fovea weakly elon-

gate, sides entirely margined, the furrows weakly crenulate, anteriorly passing

into the notauli. Prescutellar furrow crenulate. Postaxillae and lateral area

of metanotum crenulate. Propodeum densely, unevenly rugose, a broad, pen-

tagonal areola indicated. Side of prothorax posteriorly longer than above, very

fine chagreened, both furrows narrowly crenulate. Sternaulus crenulate, extend-

ing to the anterior margin, not reaching to middle coxa, epicnemial furrow

scarcely crenulate, hind marginal furrow simple. Metapleuron shining, rugose

posteriorly. Hind femur 3.5 times as long as wide, reaching near the tip of

metasoma. Wing: Stigma rather broad, r arising from basal third, rl half

as long as width of stigma, forming a straight line with r2, r2 1.66 times as

long as ccl r3 curved to outside, 1.5 times as long as r2, R reaching the tip,

nr postfurcal, Cu2 very broad, parallel sided, ccl only 1.2 times as long as cc2

d 1.7 times as long as nr, nv slightly postfurcal, B closed, twice as long as

wide, np arising from middle of B; nr in hind wing missing. Metasoma: Ter-.

gite 1 as long as wide posteriorly, evenly narrowed towards base, densely

coriaceous, dull, dorsal carinae straight, converging towards apex, gradually

merging in the sculpture. Tergite 2 fine and densely coriaceous, 3 feebly

reticulate only. Terebra hidden, the sheaths (in lateral view) shorter than

tergite 1.

224 Contrib. Amer. Ent. Inst., vol. 20, 1983

Figs. 1-3. Opius (Grimnirus) eutownesi, n. sp.: 1, Head with antennal

base in frontal view. 2, Fore and hind wing. 3, Hind leg.

Fischer: Opius (Braconidae) 225

Black. Brown: legs, especially the posterior ones and the femora pre-

dominantly, clypeus, mouth parts, anellus and wing venation. Tegulae dark.

Wing membrane feebly brownish.

Length of body: 2 mm.

Male: Antennae 28-jointed. Mesopleuron anteriorly feebly reticulate.

Variability: Antennae 25- 28-jointed.

Holotype: Female, and Paratypes: | male, 5 females, Ile-Ife, Nigeria,

May 1973, J. T. Medler (CPH, Gainesville).

Etymology: The species is names in honor of Dr. H. K. Townes.

2. Opius (Grimnirus) ileifensis, n. sp. (Figs. 4 - 6)

The species is very fine shagreened over metasomal tergite 2. Therefore

it has to be ascribed to the subgenus Grimnirus Fischer. The species closest

to it is Opius katonicus Fischer from which it is distinguished by the follow-

ing characters: cheek considerably shorter than width of mandible at base,

ocelli arranged in an equilateral triangle, clypeus 2.5 times as wide as high,

Cu2 strongly narrowed distad, delimted by straight veins, ccl 2.5 times as long

as cc2, metasomal tergite 1 1.25 times as long as apically wide and rather

shining.

Female: Smooth: head, mesoscutum, scutellum, postaxillae and the

posterior part of metasoma. Head 2.2 times as wide as long, 1.35 times as

wide as mesoscutum, 1.9 times as wide as face, 2.4 times as wide as meta~

somal tergite | at apex; laterally and on occipus with fine hairs, eyes protrud-

ing, twice as long as temples, eyes and temples commonly rounded, distance

of toruli from each other greater than distance from eyes, occiput feebly sin-

uate; ocelli strongly protruding, arranged in an equilateral triangle, distance

from each other as long as diameter of an ocellus, distance of a lateral ocel-

lus from eye as long as width of ocellar area. Face 1.3 times as wide as

high, with fine and rather long hairs, very fine and sparsely punctate, median

carina only feebly developed, eye margins parallel. Clypeus 2.5 times as wide

as high, separated from face by an evenly curved furrow, lower edge straight,

in the same plane as the face, with a row of distinct bristle points. Distance

of tentorial pits from each other three times as great as distance from an

eye. Gena considerably shorter than width of mandible basally. Mouth

closed, mandibles not expanded at base, maxillary palpi as long as the head

high. Eye in lateral veiw 1.5 times as high as long, 2.5 times as long as width

of temple, the latter parallel sided. Antennae 1.5 times as long as body, 27-

jointed; flagellar joint 1 three times, joint 12 twice, the penultimate joint 1.7

times as long as wide; hairs shorter than width of flagellar segments, in later-

al view three sensillae visible. Thorax 1.33 times as long as high, 1.3 as high

as head, upper side strongly convex. Mesoscutum 1.25 times as wide as long,

in front of the tegulae rather evenly rounded, middle lobe little protruding,

notauli only anteriorly impressed, feebly sculptured, dorsal fovea narrow and

elongate, sides only near the tegulae margined. Prescutellar furrow crenulate.

Propodeum irregular rugose. Side of prothorax posteriorly a little longer than

above, scarcely noticeably shagreened, the furrows distinct, but only very

feebly crenulate. Mesopleuron weakly shagreened, sternaulus crenulate and

partly rugose, reaching to anterior margin, ending in front of middle coxa,

226 Contrib. Amer. Ent. Inst., vol.-20, TGS.

ere

0,5: mm

Figs. 4-6. Opius (Grimnirus) ileifensis, n. sp. : 4, Head with antennal

base in dorsal view. 5, Head in ventral view. 6, Fore wing.

Figs. 7-9. Opius (Grimnirus) nigeriensis, n. sp. : 7, Head in dorsal

view. 8, Propodeum and metasoma in dorsal view. 9, Fore and hind wing.

Fischer: Opius (Braconidae) 227

the remaining furrows simple. Metapleuron shining, uneven, somewhat rugose

towards edges. Hind femur 5.5 times as long as wide, reaching to the tip of

metasoma. Wing: Stigma wedge shaped, r arising proximad of middle, r1 0.33

times as long as width of stigma, forming a straight line with r2, r2 1.3

times as long as nr, nv as long as the own width postfurcal, B closed, 2.5

times as long as wide, a little enlarged distally, np arising from middle of B;

nr in hind wing missing. Metasoma: Tergite 1 1.25 times as long as apical-

ly wide, towards base evenly narrowed, posterior width 1.5 times as long as

basal one, shining, with a longitudinally striate tendency, dorsal carinae

markedly developed only near base, converging in basal half, parallel in apical

half, reaching near hind margin. Tergite 2 very fine shagreened. Terebra

scarcely projecting, in lateral view as long as tergite 1.

Black. Yellow: Scape, anellus, clypeus, mouth parts, legs, tegulae, wing

venation and ventral side of metasoma at base. Dirty yellow: anterior half

of dorsal surface of metasoma, tergite 1 with dark shadow. Wing membrane

hyaline.

Length of body: 1.9 mm.

Male: Unknown.

Holotype: Female, Ile-Ife, Nigeria, July 1973, J. T. Medler (CPH,

Gainesville).

3. Opius (Grimnirus) nigeriensis, n. sp. (Figs. 7 -9)

The species is ascribed to the subgenus Grimnirus Fischer. The tax-

onomically nearest species is Opius bururianus Fischer from which it is dis-

tinguished by the shorter metasomal tergite 1 (apically 1.2 times as wide as

long), and the thick hind femora (4 times as wide as long).

Female: Head, thorax and metasoma apically smooth. Head 2.2 times

as wide as long, 1.3 times as wide as mesoscutum, 1.75 times as wide as face,

1.8 times as wide as metasomal tergite 1; eyes protruding, at temples strongly

narrowed, upper side only laterally and on occiput feebly hairy, distance of

toruli from each other longer than distance from eyes, occiput weakly sinuate;

ocelli large and protruding, distance from each other as long as diameter of

an ocellus, distance of lateral ocellus from eye as long as width of ocellar

area. Face as wide as high, with very fine punctures and bright hairs, median

carina narrow and without hairs, eye margins ventrad a little diverging.

Clypeus 1.7 times as wide as high, delimited by a semicircular suture, rather

flat, lower edge rounded. Tentorial pits small, distance from each other

twice as long as distance from an eye. Genae as long as basal width of man-

dible. Mouth closed, mandibles at base not expanded. maxillary palpi as long

as height of head. Eye in lateral view 1.8 times as high as long, twice as

long as width of temple, marginal furrow of temple feebly crenulate. Anten-

nae 1.33 times as long as body, 23-jointed; flagellar joint 1 three times as long

as wide, a trifle narrower than joint 2, joint 10 and the penultimate twice as

long as wide; hairs shorter than width of joints, in lateral view three sensillae

visible. Thorax 1.1 times as long as high, 1.33 times as high as head, dorsally

strongly rounded. Mesoscutum 1.3 times as wide as long, in front of tegulae

evenly rounded, notauli anteriorly deep, crenulate, reaching on disc, disappear-

ing there, dorsal fovea drop-shaped, sides margined, the marginal furrows

228 Contrib. Amer, Gat mst.) von 20, 1983

10 1. am

Fig. 10. Opius (Grimnirus) obuduensis, n. sp. bodyin lateral view.

Contrib. Amer. Ent. Inst., vol. 20, 1983 229

crenulate, anteriorly passing into notauli. Praescutellar fovea with some

carinae in the depth. Lateral areas of metanotum crenulate. Propodeum be-

hind base declivious, densely reticulate, with feebly indicated pentagonal

areola, basal carina, central carina and costulae. Side of prothorax poster-

iorly 1.8 times as long as above, both furrows crenulate, uniting below.

Sternaulus broad, oval, crenulate, on both sides shortened, epicnemial furrow

deep, sinuate, inconspicuously crenulate, posterior marginal furrow feebly

punctate, nearly simple. Metapleuron rugose. Hind femora 4.0 times as

wide as long, reaching as far as tip of metasoma. Wing: Stigma moderately

wide, r arising before middle, rl 0.4 times as long as width of stigma, form-

ing an obtuse angle with 12, r2 1.8 times as long as ccl, r3 curved outside,

1.7 times as long as r2, R reaching to wing apex, nr postfurcal, Cu2 parallel

sided, ccl 1.33 times as long as cc2, d 1.9 times as long as nr, nv 0.5 of own

length postfurcal, B distad a little widened, 3.5 times as long as wide, closed,

np arising from middle of B; nv in hind wing missing. Metasoma: Oval, nearly

as broad as thorax. Tergite 1 apically 1.2 times as wide as long, evenly nar-

rowed to base, posteriorly 1.2 times as wide as at base, dorsal carinae straight-

ly converging to hind margin, delicately granulate, basally shining. Tergites

2 and 3 densely, delicately granulate, hind margin of tergite 3 and the rest

of the metasoma shining. Terebra hidden, sheaths in lateral view shorter than

tergite 1.

Black. Yellow: clypeus, pedicel, anellus, wing venation and the ventral

surface of metasoma at base. Dark brown: legs predominantly, tegulae and

mouth parts more or less. Wing membrane hyaline.

Length of body: 1.8mm.

Male: Unknown.

Holotype: Female, Ile-Ife, Nigeria, August-September 1974, J. T. Medler

(CPH, Gainesville).

4. Opius (Grimnirus) obuduensis, n. sp. (Fig. 10)

With regard to the very fine shagreen over the middle of the gaster to

be ascribed to the subgenus Grimnirus Fischer. Because of the long thorax

closest to Opius fuscicarpus Szépligeti, and oO. toromojoensis, n. sp.

O. obuduensis, n. sp. is distinguished from them by the strongly crenulate

sternaulus, the rather thin sheaths of terebra of female, and the predominant-

ly dark coloration of body and legs.

Female: Predominantly smooth and shining. 1.8 times as wide as long,

1.2 times as wide as mesoscutum, 1.75 times as wide as face, 1.66 times as

wide as metasomal tergite | at apex; eyes not protruding, at the temples as

wide as at eyes, the latter as long as temples, only laterally and the occiput

with few fine hairs, occiput only feebly sinuate, distance between toruli as

long as their distance from eyes; ocelli scarcely protruding, small, distance

between ocelli longer than the diameter of an ocellus, distance between later=

al ocellus and eye margin as long as ocellar area wide. Face 1.2 times as

wide as high, feebly shagreened to rugose, especially near compound eyes fine

and rather densely punctuate and hairy, median carina distinctly developed and

smooth, eye margins parallel. Clypeus twice as wide as high, separated from

face by a simple, semicircular furrow, lower edge a little sinuate, situated

in the same plain as face; with some hair-bearing punctures. Distance of

230 Contrib. Amer. Ent. Inst., vol. 20, 1983

Fig. 11. Opius (Grimnirus) toromojaensis, n.sp. Lateral view.

Fischer: Opius (Braconidae) 25)

tentorial pits from each other twice as long as distance from an eye. Mouth

closed, mandibles at base not expanded, maxillary palps as long as height of

head. Genae as long as basal width of mandible. Eye in lateral view 1.4

times as high as long, as long as temple wide, the latter broader below than

above. Antennae as long as body, 27-jointed; flagellar segment 1 3.5 times,

segment 10 three times, the penultimate 1.8 times as long as wide; segments

of apical fourth gradually shorter; hairs shorter than width of segments, in

lateral view three sensillae visible. Thorax 1.5 times as long as high, 1.5

times as high as head, upper surface weakly rounded. Mesoscutum 1.15 times

as wide as long, in front of the tegulae evenly rounded, notauli anteriorly

impressed, deep and crenulate, dorsal fovea only little elongate, sides mar-

gined, anteriorly crenulate, the marginal furrows passing into the notauli.

Prescutellar furrow crenulate. Metanotum feebly rugose. Propodeum strongly

and rather densely rugose. Furrows of sides of pronotum broad, crenulate and

uniting below. Sternaulus broad, crenulate, somewhat rugose, reaching as far

as anterior margin, posteriorly shortened. Epicnemial furrow weakly crenulate,

posterior marginal furrow narrow and feebly crenulate. Metapleuron predomi-

nantly with minute rugosity. Hind femur three times as long as wide. Wing:

Stigma moderately broad, wedge shaped, r arising from before middle of stig-

ma, rl half as wide as stigma, r2 1.6 times as long as ccl, r3 straight, 1.7 times

as long as r2, R reaching tip of wing, nr postfurcal, Cu2 distad a little nar~

rowed, d 1.25 times as long as nr, nv only a little postfurcal, B posterodistad

open, d passing in a sinuate line into np; nr in hind wing missing. Metasoma:

Tergite | a trifle shorter than apically wide, only a little narrowed towards

base, dorsal carinae developed only in basal third; densely reticulate, dull, only

a little convex. Tergite (2 + 3) very feebly shagreened. The remainder of

metasoma without sculpture. Sheaths of terebra only a little projecting, in

lateral view little longer than tergite 1, hypopygium not reaching entirely as

far as tip of metasoma.

Black: head, thorax, hind coxae and metasomal tergite 1. Dark brown:

hind legs, and fore and middle legs in part; the remainder of the legs brown.

Hind margins of tergites dark. Yellow: genae, mouth parts and anelli. Anten-

nae and wing venation brown, wing membrane hyaline.

Length of body: 3 mm.

Male: Unknown.

Holotype: Female, Obudu, Nigeria, April 13, 1973, J. T. Medler (CPH,

Gainesville).

5. Opius (Grimnirus) toromojaensis, n. sp. (Fig. 11)

Among the species of the subgenus Grimnirus Fischer nearest to Opius

fuscicarpus Szepligeti. The new species is distinguished from the latter by

metasomal tergite 1, which is as long as apically wide, by the bright reddish-

brown coloration of the body, and the minor size of the body (2.8 mm).

Female: Predominantly bald, in part shining, partly shagreened. Head 1.8

times as wide as long, 1.3 times as wide as mesoscutum, 1.75 times as wide

as face, eyes only a little protruding, 1.2 times as long as temples, the latter

posteriorly narrowed, distance of toruli from each other and from eyes as long

as their diameter, occiput clearly sinuate; ocelli protruding, distance between

them as long as the diameter of an ocellus, distance between lateral ocellus

232 Contrib. Amer. Ent. Inst., vol. 20, 1983

and compound eye as long as width of ocellar area. Face 1.25 times as wide

as high, uneven, very fine and bright hairy, median carina very blunt, widened

ventrad and shining, eye margins parallel. Clypeus 2.2 times as wide as high,

separated from face by a simple, semicircular furrow, lower edge rounded, in

the same plain as the face, with fine, bright hairs. Distance between tentor-

ial pits twice as long as distance from an eye. Genae as long as width of

mandibles at base. Mouth closed, mandibles at base not expanded, maxillary

palpi as long as height of head. Eye in lateral view 1.5 times as high as long,

1.1 times as long as width of temple, the latter parallel sided. Antennae of

the present example damaged, 25 joints visible, probably longer. than body;

flagellar segment | three times as long as wide, segment 2 as long as segment

1, the last visible segment twice as long as wide; hairs shorter than antennal

segments wide, in lateral view 2 or 3 sensillae visible. Thorax 1.5 times as

long as high, 1.2 times as high as the head, dorsal surface only weakly convex,

nearly parallel with the ventral surface. Mesoscutum as wide as long, lateral

lobes rounded, middle lobe little protruding, notauli anteriorly deep and crenu-

late, reaching on disc, but here disappearing, dorsal fovea short, narrow, mar-

gined on lateral edges, the marginal furrows passing into the notauli. Pre-

scutellar furrow crenulate. Lateral areas of metanotum feebly striate. Pro-_

podeum densely, fine rugose, dull. Side of prothorax posteriorly as long as

above, very fine shagreened, furrows without crenulation. Mesopleuron very

fine shagreened, sternaulus impressed, only shagreened, posterior marginal fur-

row simple. Metapleuron partly shagreened and with long, bright hairs. Hind

femora three times as long as wide, not reaching as far as tip of metasoma.

Wing: Stigma moderately wide, wedge shaped, r arising a little from before

middle, rl 0.25 times as long as width of stigma, forming a blunt angle with

r2, r2 1.66 times as long as ccl, r3 nearly straight, twice as long as r2, R

reaching tip of wing, nr postfurcal, Cu2 narrowed distad, d 1.33 times as long

as ccl, nv as long as the own width postfurcal, B posterodistad open, d passing

in a sinuate line into np; nv in hind wing only feebly indicated as a fold.

Metasoma as long as head and thorax together. Tergite | as long as apically

wide, evenly narrowed to base, dorsal carinae developed only at base and far

distant from each other, fine and densely granulate, at hind corners shining.

Tergites 2 + 3 feebly shagreened, the sculpture disappearing posteriorly. The

remainder of metasoma shining. Sheaths of terebra little projecting, in later-

al view 1.7 times as long as Tergite 1. Tip of hypopygium incised, hypopygium

ending considerably before tip of metasoma.

Bright reddish brown. Dark: flagellum, tips of mandibles, apical tarsal

segments and sheaths of terebra. Yellow: mouth parts, tegulae and wing

venation. Wing membrane hyaline.

Length of body: 2.8 mm.

Male: Unknown.

Holotype: Female, Toromoja, Botletle, Botswana, April 26, 1975, Peter

Ginn (CPH, Gainesville).

Subgenus FREKIUS Fischer

Subgenus Frekius Fischer, 1971. Ann. Naturh. Mus. Wien 75: 390.

Type-species: Opiuscastaneus Fischer (designated by Fischer 1972).

Appeared actually 1972. Fischer, 1972. Das Tierreich 91: 70, 75 (description).

Fischer: Opius (Braconidae) 233

Fischer, 1977. Das Tierreich 96: 67 (New World species).

Mesoscutum with dorsal fovea, precoxal suture sculptured, mouth closed,

mandibles not expanded at base, clypeus without tooth on lower edge, head

dorsally smooth or at most occiput punctuate, metasoma beyond tergite |

smooth.

There are 5 New World and 11 Old World species. One African species

is described as new in the present paper.

10.

KEY TO THE OLD WORLD SPECIES

fe ANCGTICa Sr IROL STIS a ee ee ae oe A cae dae ee 2

WEY “OMT EAE a yh ee ew ae aN Sar ek eo Ge ee | eal ast 6

Stigma wide, hall .ege<snaped 05/4. c\e eet ke RR aw wee woke 3

Stigma narrow, wedge-shaped, or if rather wide, then triangular ... 5

Mesoscutum evenly and intensively hairy. 3.2 mm. M. Canary Islands .

CT aT Vee eee ee Cera em ew ee ES beckeri Fischer

Hairy only declivity of mesoscutum, two stripes along the imaginary

course of notauli, and the lateral margins anteriorly more or less . 4

Body nearly entirely brown. Epicnemial furrow and anterior mesosternal

furrow crenulate. Hind femora 4 times as long as wide. Median area

of metasomal tergite 1 evenly rugose. Base and tip of stigma pale.

3.8 mm. F. USSR (Guzar, Uzbekistan) ..... imitabilis Telenga

Black: Head, thorax predominantly, and metasomal tergite |. Epicnemial

furrow and anterior mesosternal furrow simple. Hind femora three

times as long as wide. Median area of metasomal tergite | only with

traces of rugae. Stigma entirely brown. 3.8 mm. F. Algeria

COBAE ou ie Sidslke de de Geta) ghana aah es ee barbieri Fischer

m2 1.25 times ‘as tone as cel. FO -mm. OMe Madneaecae wy ok wre

RE WN. Rey ONE yO er Ae, nO ee om ae ee ns, a contrarius Fischer

r2 1.5 times as long as ccl. 2.6 mm. M. Zaire. . fukuensis Fischer

Head less than twice as wide as long, not narrowed beyond eyes ... 7

read. at lasercwWiee 2s-Wide: a6 AONE: 5. vowel asl ad ee nee bathe wet aa 9

Thorax 1.6 times as long as high. Posterior marginal furrow of meso-

pleuron and epicnemial furrow simple. 6.0 mm. M. Madagascar:

Ce a he el Re ae eat aeagd Mite Mem coed re, eee cf. seyrigi Fischer

Thorax 1.33 times as long as high. Posterior marginal furrow and

epicnemial furrow crenulate. 2.2 mm. M. USSR (Kazan, Tatarskaya

ABS yy ry A mee eno) irae he nan eae ot rossicus Szépligeti

Thorax 1.5 times as long as high. 2.6 mm. M, F. Madagascar... .

. subangulatus Granger

Thorax Ldoote, Lae: Wied a9 Hone Oe Nitin, oe al eles Wie Ane e aek wE 9

R ending clearly before tip of wing. F. 1.8 mm. Bulgaria .......

iy a MR ae as Oleg ae Cd a Rear cf. plovdivensis Zaykov & Fischer

Pe POAC TER Wee SE em i es ey a ne nO ne oak eae ceae 10

Thorax reddish brown. Projecting part of terebra 0.33 times as long as

metasoma. 2.7 mm. F. Queensland . .. terraereginae Fischer

Fig. 12. Opius (Frekius) nairobicus,n.sp. Lateral view.

Fischer: Opius (Braconidae) | 235

Thorax entirely or predominantly black. Terebra either scarcely project-

ing or at least 0.75 times .as long as metasoma. 0.0.5 was 402 11

11. Terebra of female scarcely projecting. 2.3 mm. F. Madagascar... .

A ie are dg Mr Me ie Ge ak a iba us ear ast ae .castaneus Granger

Terebra of female at least 0.75 times as long as metasoma...... 12

12. Legs entirely yellow. Terebra something longer than metasoma. Head

2.3 times. as wide as, long... 4.1.mm...8. | KeNIAs io 62 is epee oe #

ee en. epee im arene gaa 1 PENNA Res aie SFA pe 1. Bazrobicus,. A. Sp.

Legs brown. Terebra 0.75 times as long as metasoma. Head twice as

wide as. lone. 4.0.mmwe, ©. , Zaire §<.- -icuae- a fubogenus.. Fischer

1. Opius (Frekius) nairobicus, n. sp. (Fig. 12)

Because of the stout thorax, the black coloration of the body, and

the long terebra among the species of the subgenus Frekius Fischer nearest

to O. lukuganus Fischer. The new species is distinguished from the latter

by the yellow legs, the broader head, and the still longer terebra of female.

Female: Smooth: the entire body except the propodeum and metasomal

tergite 1. Head 2.3 times as wide as long, 1.15 times as wide as mesoscutum,

1.8 times as wide as face, twice as wide as metasomal tergite | apically; dor-

sal surface only with inconspicuous hairs; eyes protruding, 2.4 times as long

as temples, eyes and temples commonly rounded, distance between toruli and

between toruli and eyes as long as diameter of toruli, occiput medially dis-

tinctly sinuate; ocelli protruding, distance between them as long as diameter

of an ocellus, distance of lateral ocellus to eye as long as width of ocellar

area. Face 1.2 times as wide as high, fine, bright and sparsely hairy-

punctured, median carina narrow, smooth and shining, eye margins parallel.

Clypeus 2.4 times as wide as high, only little convex, separated from face by

a little curved suture, lower edge rounded, with very fine hairs and punctures.

Genae shorter than basal width of mandible. Mouth closed, mandibles not ex~=

panded at base, maxillary palpi as long as height of head. Eye in lateral

view twice as high as long, 1.25 times as long as width of temple, parallel

sided. Antennae 1.5 times as long as body, 34-jointed; flagellar segment |

three times, segment 12 2.2 times, and the penultimate twice as long as wide;

hairs as long as width of antennal segments, in lateral view 4 sensillae visible.

Thorax 1.3 times as long as high, 1.6 times as high as head, dorsal surface

strongly convex. Mesoscutum 1.2 times as wide as long, in front of tegulae

rather evenly rounded, middle lobe only a little protruding; notauli deeply im-

pressed anteriorly, inconspicuously sculptured, reaching on disc, but there

disappearing, the imaginary course indicated by a band of fine hairs, dorsal

fovea distinct, sides only near tegular margined. Prescutellar furrow with some

longitudinal ribs. Postaxillae medially crenulate. Lateral areas of metanotum

with few inconspicuous crenulae. Propodeum densely reticulate. Side of pro-

thorax posteriorly longer than above, the furrows crenulate, uniting below.

Sternaulus broad, oval, irregularly crenulate and rugose, the other furrows

simple. Metapleuron rugose, shining in the middle. Hind femora 5 times as

long as wide, not entirely reaching as far as tip of metasoma. Wing: Stigma

wide, wedge-shaped, r arising from basal third, rl 0.4 times as long as width

of stigma, passing into r2 in a sinuate line, r2 1.5 times as long as ccl, r3

236 Contrib. Amer. Ent. Inst., vol. 20, 1983

curved outward, 1.5 times as long as r2, R reaching tip of wing, nr strongly

postfurcal, Cu2 distad strongly narrowed, ccl weakly bisinuate and 2.35 times

as long as cc2, d twice as long as nr, nv little postfurcal, B closed, 2.1 times

as long as wide, distad only little widened, np arising from middle of B; nr

in hind wing missing. Metasoma: Tergite | as long as apically wide, poster-

iorly with parallel sides, basad little narrowed, dorsal carinae near base con-

verging, parallel from middle to hind margin, median area only a little ele-

vated, rugose, lateral areas shining. Sheaths of terebra as long as metasoma

and thorax together. Hypopygium reaching near tip of metasoma.

Black. Yellow: base of antennae in part, clypeus, genae, mouth parts,

all legs, tegulae, wing venation and a part of the ventral side of metasoma.

In strong light two brown stripes and a pitch on disc of mesoscutum visible;

the same the anterior and posterior edges of metanotum. Wing membrane hy-

aline.

Length of body: 3.1 mm.

Male: Unknown. |

Holotype: Female, Karen, Nairobi, Kenya, October 1972, 2000+ m.

C'ngaham -— Van Someren. Paratype: 1 female, same date, December 1972

(CPH, Gainesville).

REFERENCES

Fischer, M. 1972. Hymenoptera, Braconidae, Opiinae I. Das Tierreich 91:

bg X13 66205

Fischer, M. 1973. Die athiopischen Opius—Arten der Sektion A, aufgeteilt

auf die Untergattungen. Ann. Naturh. Mus. Wien 75: 387-433.

Fischer, M. 1973. Uber die afrikanischen Opiinae der Sammlung Townes in

American Entomological Institute in Ann Arbor, Michigan. Polskie Pismo

Ent. 43: 241-272.

Fischer, M. 1977. Hymenoptera, Braconidae, Opiinae II ~ Amerika. Das

Tierreich 96: I-XXVII, 1-l001.

Fischer, M. _ Illustrierte Redeskriptionen von Opiinen aus der UdSSR und neuer

Bestimmungsschlussel zu den palaarktischen Arten des Subgenus Utetes

Foerster. Ann. Naturh. Mus. Wien, in press (0. imitabilis Telenga

ascribed to subgenus Frekius Fischer).

Zaykov, A. and Fischer, M. Uber Opiinae aus Bulgarien. Ann. Naturh. Mus.

Wien, in press (description of 0. plovdivensis, n. sp.).

THREE NEW OPIINAE SPECIES FROM INDIA

(HYMENOPTERA : BRACONIDAE)

J. Papp

Zoological Department, Hungarian Natural History Museum

Budapest, Hungary

Abstract

Three new Opiine species from India are described: Biosteres townesi,

n. sp., Opius marjorieae, n. sp. and Opiustownesianus , n. sp., with 12 figures.

Dr. G. Topal, Mammoligist of the Hungarian Natural History Museum,

Budapest, had been on two collecting trips in India during 1966-1967 and

1979 -1980 for one year each. He had collected animals for the museum and

among this material there are about 150-200 specimens of Braconid wasps.

From among the Opiinae four specimens proved to represent three new species:

Biosteres (Biosteres) townesi, n. sp, Opius (Apodesmia) marjorieae, Nn. sp.

and Opius (Utetes) townesianus, n. sp. They are described below and the

specific differences are tabulated indicating the nearest related species. The

descriptions are accompanied with 12 original figures.

1. Biosteres (Biosteres) townesi, n. sp. (Figs. | -4)

Male: Body 3.2 mm ong. Head in dorsal view almost twice broader than

long, temple 1.7 times longer than eye, head behind eyes distinctly broadening,

i.e., head between templed clearly broader than between eyes (Fig. 1). Toruli

slightly nearer to each other than to eye. Face together with clypeus one-

third wider than high. Clypeus thrice wider below than high medially. Distance

between tentorial pits almost twice greater than distance between a pit and

lowest point of eye. Cheek as long as basal width of mandible. Mouth closed.

Mandible gradually broadening basally, below its basal half with a strong rim

ending somewhat angularly. Eye in lateral view twice higher than wide, temple

distinctly 1.5 times wider than eye (Fig. 2). Ocelli rather small, distance be-

tween two ocelli equal with greatest diameter-of elliptic hind ocellus. OOL

almost thrice as long as P9L. Head polished, face and clypeus with shallow

hair punctures. Antenna somewhat longer than body, with 35 joints. First

flagellar joint thrice as long as broad, further joints slightly shortening and

attenuating so that penultimate joint also thrice as long as broad.

Mesosoma in lateral view 1.4 times longer than high. Mesonotum as long

medially as wide between tegulae. Notaulix restricted to declivous part of

mesonotum and here shallow to very shallow, on disc of mesonotum notaulix

indistinct. Mesonotal dimple linear and deep. Prescutellar furrow with weak

crenulation. Postaxille smooth. Propodeum smooth, its lateral spiracle some-

what protruding. Sternaulix indicated by a very feeble linear impression.

Hind femur 3.6 times as long as broad. Hind tibia and tarsus equal in length.

Fore wing somewhat longer than body. Stigma (Fig. 4) linear in form,

i.e., its sides parallel and issuing radial vein from its proximal half. rl about

as long as width of stigma, r2 somewhat shorter than cuqul, r3 thrice as long

as r2 and ending before tip of wing. Cu2 narrowing distally, n. rec. post-

furcal (Fig. 4). d distinctly 1.4 times longer than n. rec., B. closed.

Metasoma about as long as head + mesosoma together. First tergite (Fig.

3) somewhat longer than wide at hind, twice wider behind than basally, with

Zul

238 Contrib. Amer. Ent. Inst., vol. 20, 1983

La a ay

SIRS IOES

Figs. 1-4. Biosteres (Biosteres) townesi, n. sp. Male: 1, Head in

dorsal view. 2, Head in lateral view. 3, First tergite. 4, Distal part of

right fore wing. Figs. 5-7. Opius (Apodesmia) marjorieae, n. sp. Female:

5, Head in dorsal view. 6, First tergite. 7, Distal part of right wing.

Figs. 8-10. Opius (Utetes) townesianus, n. sp. M, F : 8, Head in dorsal view

behind eyes. 9, Distal part of right fore wing. 10, First tergite. Figs. 11-12.

Opius (Utetes) infernalis Fischer: 11, Head in dorsal view. 12, First tergite.

Papp: Opiinae (Braconidae) 239

a pair of converging keel on its basal half. Surface of hind half of first

tergite shiny and with very fine longitudinal rugulo-striation. Further tergites

polished.

Head and metasoma dark brown, mesosoma brownish yellow. Clypeus, man-

dible and temple below brownish yellow, flagellum blackish brown, palpi pale.

Mesopleuron, metanotum and propodeum dark brown to brown. Second tergite

brownish yellow. Legs brownish yellow, hind tibia and tarsus rather yellow.

Wings hyaline. Stigma and venation brownish yellow.

Female and host: Unknown.

Holotype: Male, India: West Bengal, Senchal Reserve Forest, Ghum, 2200

m, 22 October 1967, leg. Topal, No. 886.

Holotype is deposited in Hungarian Natural History Museum, Budapest; Hym.

Typ. No. 5312.

The new species is dedicated to Dr. Henry Townes.

From among the Indo-Australian Biosteres species the new species seems

closely allied with Biosteres (B.) kashmirensis (Fischer 1966). They may

be distinguished by the following features:

B. townesi B. kashmirensis

1. Head in dorsal view behind eyes 1. Head in dorsal view behind eyes not

broadening, i.e., head between not broadening, i.e., head between

temples distinctly broader than eyes and temples about equal breadth.

between eyes (Fig. 1).

2. Temple in dorsal view 1.7 times 2. Temple and eye in dorsal view of

longer than eye (Fig. 2). equal length.

3. Antenna with 35 joints, first 3. Antenna with 45 joints, first flagel-

flagellar joint thrice, penultimate lar joint 2.5 times and penultimate

joint also thrice longer than broad. joint 1.5 times longer than broad.

4. R ending before tip of wing (Fig. 4. R reaching tip of wing.

4).

5. Propodeum smooth, shiny. 5. Propodeum rugulose, dull.

6. Head and metasoma dark brown, 6. Head and mesosoma black, metasoma

mesosoma brownish yellow. reddish brown.

2. Opius (Apodesmia) marjorieae, n. sp. (Figs. 5 -7)

Female: Body rather stout, 2.2 mm long. Head in dorsal view transverse,

2.1 times broader than long, eye 2.5 times longer than temple, head behind

eyes strongly rounded (Fig. 5). Toruli nearer to eye than to each other.

Face together with clypeus indistinctly wider than high. Clypeus 2.5 times

wider below than high medially. Distance between tentorial pits twice greater

than distance between a pit and lowest point of eye. Cheek somewhat shorter

than basal width of mandible. Mouth open. Mandible gradually broadening

basally, i.e., without subtooth at its lower base. Eye in lateral view 1.9

times higher than wide, eye twice wider than greatest width of temple.

Ocelli average in size; distance between fore and a hind ocelli equal with

240 Contrib. Amer. Ent.:Inst,,. yol. 20, 1983

greatest diameter of elliptic hind ocellus; distance between hind two ocelli

somwwhat shorter than diameter indicated. OOL twice as long as POL. Head

polished. Antenna about as long as body, with 23 joints. First flagellar joint

twice, penultimate joint 1.5 times as long as broad, flagellum distally not at-

tenuating.

Mesosoma in lateral view only somewhat longer than high. Mesonotum

one-fifth wider between tegulae than long medially. Notaulix restricted to

declivous part of mesonotum, deep, on disc of mesonotum absent. Mesonotal

dimple round, deep. Postaxille entirely smooth, shiny. Propodeum rugose.

Sternaulix short, crenulated. Hind femur four times longer than broad. Hind

tarsus somewhat shorter than hind tibia.

Fore wing somewhat longer than body. Stigma not wide, four times as

long as wide, issuing radial vein close before its middle (Fig. 7); rl short, 12

1.7 times longer than cuqul, r3 somewhat longer than r2 and reaching tip of

wing. Cu2 narrowing distally, n. rec. postfurcal (Fig. 7). d twice as long as

n. rec., B closed.

Metasoma somewhat longer than mesosoma. First tergite (Fig. 6) slight-

ly wider at hind than long medially, evenly broadening posteriorly; basal con-

verging pair of keel relatively short and merging into median and rather

longitudinal rugosity, lateral surface almost smooth to smooth, shiny. Second

tergite with a pair of posteriorly diverging furrow at its fore middle, other-

wise together with further tergites polished. Tergites 2~3 fused. Ovipositor

sheath in lateral view very short, as long as second joint of hind tarsus.

Body black, legs yellow. Clypeus, labrum and mandible reddish yellow,

palpi and galea yellow. Scape and pedicel yellow, flagellum blackish. Tegula

brown, parategula brownish yellow. Middle femur faintly brownish; hind femur

brown, basally and apically yellow. Last joint of tarsi 2-3 brownish, claws

and pulvilli blackish. Wings hyaline, stigma and veins with brown pigmentation.

Male and host: Unknown.

Holotype: Female, India: Karnataka, Balgaum district, Talewadi, 780 m,

netting in grasses, 26 February 1980, leg. Topal, No. 135.

Holotype deposited in the Hungarian Natural History Museum, Budapest;

Hym. Typ. No. 5313.

The new species is dedicated to Mrs. Marjorie Townes.

Within the subgenus Apodesmia Foerster so far two species were reported

from the Indo-Australian Region. The new species, 0. marjorieae may be

separated from them as in the key below:

1. First tergite 1.8 times longer than broad at hind. Fore wing hyaline, be-

low stigma with a brown streak. Head and mesosoma densely haired.

Propodeum rugo-rugulose, with a pair of lateral smooth field. In

dorsal view eye one-fifth longer than temple. Antenna with 42-43

fOInts, “42-4 Gm, Taiwan oo maculipennis Enderlein

First tergite as long as wide at hind or slightly wider behind than long

(Fig. 6). Fore wing hyaline, without brown streak below stigma.

Head and mesosoma not densely haired. Propodeum evenly rugose,

without lateral smooth field. In dorsal view eye at least twice as

long as temple. Antenna at most with 30 joints ........... 2

Papp: Opiinae (Braconidae) 241

2. Second tergite polished. r2 1.7 times longer than cuqul, r3 only somewhat

longer than r2, stigma less wide (Fig. 7). Antenna with 23 joints, first

flagellar joint twice, penultimate joint 1.5 times as long as broad.

Postaxille smooth and shiny. Scape and pedicel yellow. Body black,

legs. Yellows hind temur brown, 2.2 mm. FF. inde eee . oe,

PA ee Ln i aca Mia err aan Siem ot geile Bee Air te. el ai, ani pec taig’ dee MOL Lea Gs Th: ais

Second tergite longitudinally striated. 12 1.25 times longer than cuqul,

r3_ 2.5 times longer than 12, stigma wide (Abb. 4, in Fischer 1978: 375).

Antenna with 30-32 joints, first flagellar joint 2.7 times, penultimate

joint 1.66 times as long as broad. Inner margin of postaxille crenulate.

Scape and pedicel black. Body reddish brown, tegulae and legs yellow.

pe 1 Ale | ae es ae oeeieenrne: pear tT alan oat formosigaster Fischer

3. Opius (Utetes) townesianus, n. sp. (Figs. 8 -10)

Female: Body 3.2 mm long. Head in dorsal view 1.8 -1.9 times broader

than long, eye twice as long as temple, head behind eyes contracted (Fig. 8).

Face together with clypeus slightly wider than high medially. Clypeus twice

wider below than high medially. Distance between tentorial pits twice great-

er than distance between a pit and lowest point of eye. Mouth open. Man-

dible gradually broadening basally, i.e., without subtooth at its lower base.

Eye in lateral view 1.4 times higher than wide, eye almost twice wider than

greatest width of temple. Ocelli relatively large and elliptic in form, distance

between fore and a hind ocelli somewhat shorter than greatest diameter of

hind ocellus. OOL nearly thrice as long as POL. Head polished, clypeus punc-

tate. Antenna one-quarter longer than body, with 36 joints. First flagellar

joint 2.5 times as long as broad, further joints gradually shortening and at-

tenuating so that penultimate joint twice as long as broad.

Mesosoma in lateral view 1.3 -1.4 times longer than high. Thorax polished.

Mesonotum as long medially as wide between tegulae. Propodeum with a

medio-longitudinal carina, medially rugo-rugulose, laterally almost smooth to

smooth. Notaulix restricted to declivous part of mesonotum. Mesonotal dimple

before prescutellar furrow round and deep. Sternaulix rather narrow and mod-

erately crenulated. Hind femur distinctly four times longer than broad. Hind

tibia and tarsus equal in length.

Fore wing as long as body. Stigma (Fig. 9) 3.5 times as long as wide,

emitting radial vein from its middle. ri short, r2 twice as long as cuqul,

r3 one-fifth longer than 12 and reaching tip of wing. Cu2 narrowing distally,

n. rec. distinctly postfurcal (Fig. 9). d almost twice as long as n. rec., B

closed.

Metasoma in lateral view slightly longer than mesosoma. First tergite

(Fig. 10) distinctly 1.5 times longer than wide at hind, moderately broadening

posteriorly, before its hind end strongly broadening; pair of keel converging

at its fore third, its further section subparallel. Surface between keels and

of hind third rugo-rugulose, otherwise first tergite smooth, shiny. Further

tergites polished. Ovipositor sheath in lateral view short, half as long as hind

tibia.

Ground color of body black, legs yellow. Clypeus, labrum and mandible

brownish yellow, palpi yellow. Surface around toruli rusty. Scape and

pedicel yellow, flagellum black. Tegulae yellow, parategulae pale yellow.

242 Contrib. Amer. Ent. Inst., vol. 20, 1983

Coxae whitish yellow, distal half of middle tibia and entire hind tibia dark

fumous, claws and pulvilli blackish. Tergites 3-6 with rusty transverse bands,

tergites 7-8 yellow. Sternites brown to light brown. Wings hyaline. Stigma

with blackish, venation with blackish brown pigmentation.

Male: Similar to the female. Body 3.3 mm long. Antenna with 34

joints. Legs more whitish yellow. Hind tarsus black.

Holotype: Female, and allotype: | male, India: West Bengal, Darjeel-

ing district, Debrapani, 1700 m, netted in grass vegetation, 29 May 1980, leg.

Topal, No. 427.

Holotype and allotype are deposited in the Hungarian Natural History Mu-

seum, Budapest; Hym. Trp. No. 5314 (holotype) and 5315 (allotype).

The new species is dedicated to Dr. Henry Townes.

The new species, 0. townesianus, n. sp., seems closely related to oO.

sanguanus Fischer, 1966 (Nepal), the specific differences between them are

few and difficult to recognize. They are tabulated below:

O. townesianus O. sanguanus

1. Body 3.2 -3.3 mm long. 1. Body 2.1 mm long.

2. Antenna with 34-36 joints. 2. Antenna with 23 joints. p)

3. Mesonotum as wide as long. 3. Mesonotum somewhat though dis-

tinctly wider than long.

4. First tergite moderately broadening 4. First tergite evenly broadening

posteriorly, and strongly broadening posteriorly (Abb. 66, in Fischer

close before its hind end. 1966: 106).

5. r3 only one-fifth as long as r2 5. r3 1.5 times as long as r2 (Abb.

(Fig...9). 64, l.c.).

6. Temple in dorsal view distinctly 6. Temple in dorsal view somewhat

half as long as eye (Fig. 8). shorter than eye.

O. townesianus is close to O. infernalis Fischer, 1966 (Philippines,

Taiwan). They differ as follows:

O. townesianus : O. infernalis

1. Head in dorsal view behind eyes 1. Head in dorsal view behind eyes

rather constricted (Fig. 8). rounded (Fig. ll).

2. Ovipositor sheath in lateral view 2. Ovipositor sheath in lateral view

short, half as long as hind tibia. long, one-sixth longer than hind tibia.

3. First tergite moderately broadening 3. First tergite rather evenly broaden-

posteriorly and strongly broadening ing posteriorly (Fig. 12).

close before its hind end (Fig. 10).

4. Ground color of head and mesosoma 4. Ground color of head and thorax

black, head around toruli rusty reddish to brownish yellow with

(M) or reddish yellow (F). more or less black pattern.

Papp: Opiinae (Braconidae) 243

Considering the somewhat elongated mesosoma of 0. townesianus, it runs

to O. nepalensis Fischer, 1966 (Nepal) in Fischer's key for the Indo-Australian

species of the genus oe These two species are separated below:

O. townesianus O. nepalensis

1. Mesosoma in lateral view 1.3 -1.4 1. Mesosoma in lateral view distinctly

times longer than high. 1.5 times longer than high.

2. Body 3.2 -3.3 mm long. 2. Body 1.7 mm long.

3. Stigma issuing radial vein from its 3. Stigma issuing radial vein from its

middle (Fig. 9). basal third.

4. Antenna 34-36 jointed. Penulti- 4. Antenna 19-21 jointed, usually with

mate joint of antenna twice as long 20 joints. Penultimate joint of

as broad. antenna thrice as long as broad.

5. Entire flagellum blackish. 5. Last 4-6 joints of flagellum whitish,

otherwise flagellum blackish.

Fischer (1980) recently described Opius (Utetes) mudigerensis from South

India, which also seems allied to townesianus. They differ as follows:

O. townesianus O. mudigerensis

l. Body 3.2 =3.3 mm long (M, F). 1. Body 4 mm long (F).

2. First tergite distinctly 1.5 times 2. First tergite 1.25 times longer than

longer than wide at hind (Fig. 10) wide at hind (Abb. 3, in Fischer

1980: 175).

3. Ovipositor sheath short, in lateral 3. Ovipositor sheath long, in lateral

view half as long as hind tibia. view as long as hind tibia.

4. Basal flagellar joints relatively 4. Basal flagellar joints relatively

long, first joint 2.5 times as long short, first joint twice as long as

as broad. broad.

5. Mesonotum as long pen as wide 5. Mesonotum 1.2 times wider between

between tegulae. tegulae than long medially (Abb.

1 Peo.

REFERENCES

Enderlein, G. 1912. H. Sauter's Formosa-Ausbeute. Braconidae, Prototrupidae

und Evaniidae (Hym.). Entomol. Mitt. 1(9): 257-267.

Fischer, M. 1966. Revisioh der Indo-Australischen Opiinae (Hymenoptera, Bra-

conidae). Ser. Entom. |: I-VI = 1-167.

Fischer, M. 1967. Redeskriptionen von einigen Opiinen (Hymenoptera,

Braconidae. Zschr. Arb. gem. Osterr. Entom. 19(2-3): 59-69.

Fischer, M. 197 1a. Opiinae aus Neu-—Guinea und von den Bismarck-Inseln

(Hymenoptera: Braconidae). Pacific Ins. 13(3-4): 487-512.

Fischer, M. 197lb. Die Opiinae der Noona Dan Expedition nach den Philip-

pinen und Bismarck-Inseln und Redeskription von pl a dissitus aus Hawaii

(Insecta, Hymenoptera, Braconidae). Steenstrupia 2: 1-25.

244 Contrib. Amer. Ent. Inst., vol. 20, 1983

Fischer, M. 1975. Drei orientalische Opiinen (Hymenoptera, Braconidae)

aus der Sammlung Townes. Zschr. Arb. Gem. Osterr. Entom. 27(1-2):

47-52.

Fischer, M. 1978. Neue Opiinae (Hymenoptera, Braconidae) von der

australischen Region, besonder aus Tasmanien. Polskie Pismo Entom. 48:

371-412.

Fischer, M. 1980. Opius mudigerensis n. sp., eine neue Braconide aus

Indien (Hymenoptera, Braconidae, Opiinae). Steestrupia 6(ll): 173-176.

Papp, J. 1982. Taxonomical and faunistical novelties of the Opiinae in the

Arctogaea (Hymenoptera, Braconidae). Ann. Hist. Nat. Mus. Nat. Hung.

74: 241-253.

STUDIES ON THE GENUS PROSTHOPORUS

(HYMENOPTERA : ICHNEUMONIDAE)

Santosh Gupta

Department of Entomology & Nematology

University of Florida, Gainesville, FL. 32611

Prosthoporus was described by Porter (1976) by a single female from

Bolivia. Additional specimens of this genus in the Townes Collection revealed

several undescribed species from South America. They are described below.

The taxonomic placement of the genus is also discussed.

Genus PROSTHOPORUS

Prosthoporus Porter, 1976. Psyche, 83: 271-276. Type-species:

Prosthoporus terani Porter. Monotypic and original designation.

Fore wing 4 to 12 mm long. Body slender. Frons without a median horn

or carina, smooth and shiny or weakly shagreened to finely striate. Subapical

antennal segments flattened. Clypeus moderately large, its basal half convex

and apical half flattened, its apical margin concave without a median tooth.

Mandible 2.2x as long as its basal width; its lower tooth strong, slightly shorter

than the upper tooth or nearly equal. Temple flat. Occipital carina touching

or not touching the hypostomal carina. Pronotum finely striate to smooth.

Epomia present though weak. Sternaulus very weak and shallow, hardly reach-

ing 0.5 its length. Prepectal carina usually less than 0.6 its length. Hind mar-

gin of metanotum without any projection on each side of metascutellum. Area

between metascutellum and propodeum long, gently sloping and shagreened.

Propodeum elongate, gently and evenly arched in profile view. Propodeal

spiracle oval to elongate. Pleural carina absent. Basal transverse carina

straight and sharp. Apical transverse carina completely absent, apophysis or

crest absent. Propodeum transversely striate, basal area shagreened to granu-

lose. Areolet of moderate size, rectangular, about 0.6 as high as section of

second recurrent vein above bulla, about 1.7 as wide as high, receiving second

recurrent vein before or after the middle. Second intercubitus weak and hard-

ly visible medially. Ramellus absent. Nervulus opposite or basad of basal vein

by about 0.25 its length. Nervellus intercepted at its apical 0.4 or at the

middle. Brachiella absent. Fore tibia in female moderately swollen and with

some usually strong and sparse, spine-like setae in addition to normal dense

hairs. Fore tarsus in female with fourth segment deeply to weakly bilobed

apically. Abdomen shagreened. First tergite moderately long and slender,

its spiracle at its basal 0.4, without any lateral tooth at base, without longi-

tudinal carinae, its sternite about 0.3 its length. Second tergite basally with

or without a few punctures. Ovipositor slender, moderately compressed, of

variable length. Ventral valve of ovipositor with strong, oblique to vertical

ridges and with a dorsal lobe that partially encloses the upper valve.

Geographical distribution and host associations: Prosthoporus was

so far known from Bolivia. It is now recorded from Brazil, Ecuador, Panama

and Peru in Central and South America. The hosts are unknown.

246 Contrib. Amer. Ent. Inst., vol. 20, 1983

AFFINITIES OF PROSTHOPORUS

Prosthoporus was described under the subtribe Lymeonina. No doubt

it shares some of the characters of the Lymeonine genera, like the lower

tooth of mandible shorter than the upper, and the fourth tarsal segment of

the female fore tarsus usually deeply bilobed. However, these characters

are also present in some other species of Gabuniini, like Pharzites nigri-

tarsis, Apocryptus terebratus, A. chinensis and Schreineria taiwana having

the lower tooth of mandible equal to or shorter than the upper, and

Xoridesopus maculatus, X. striatus and species of Cestrus having the fourth

segment of female fore tarsus bilobed. Prosthoporus appears more related

to the Gabuniini, particularly by the following combination of characters:

Spiracle of first tergite at its basal 0.4, tip of lower valve of ovipositor with

a dorsal lobe that partly encloses the upper valve, shape of the areolet, in-

flated fore tibia of female, propodeal apophyses usually absent, and first ter-

gite without median dorsal carinae. Therefore the genus Prosthoporus is

transferred to the Gabuniini.

Within the Gabuniini, the genus Prosthoporus shows affinities with

Gabunia, Torbda and Agonocryptus. It is distinguished from Gabunia by the

rectangular areolet, from Torbda by the absence of the transverse ridge and

apical tooth on the clypeus, and from Agonocryptus by its weak sternaulus,

shorter lower mandibular tooth, absence of brachiella, absence of apical trans-

verse carina of propodeum, sculpture of propodeum, and by its long excavated

and shagreened area between the metanotum and propodeum.

KEY TO THE SPECIES OF PROSTHOPORUS

1. Thorax and abdomen reddish-brown, with white marks and bands. Pro-

notal groove finely striate. Frons granulose or finely striate. Malar

space 0.5 or more its basal width. Length of ovipositor equal to the

tener of Wied Iie ty wk Gay che aries ee tee A ewes Soaiondeus 2

Thorax brownish-red with abdomen black, with yellow bands. Pro-

notal groove smooth and shiny. Frons usually smooth. Malar space

0.25 to 0.4 its basal width. Ovipositor much longer than the length

efonind tiple (equal Imshs berant sau och Geichaws bs race s ee 3

2. Frons granulose. Wings banded. Face as wide as high. Brazil ......

Se tor Ue eine ee ere eo a Game te mec ee eer UR Gn i. townesi, Nn. Sp:

Frons finely striate. Wings not banded. Face wider than high. Brazil .

Sah Gd es ee a ee a eee ie ee 2. Lriacirrons) “Nn. So:

3. Clypeus shiny. Mandible with fine punctures. Yellow orbital ring com-

plete. Hind femur yellow. Trochanters of hind leg yellow. 4th

tarsal isepment of hind jes Vellow:. 2. & on wad ede we era ~ we he 4

Clypeus dull. Mandible mat. Yellow orbital ring incomplete. Hind femur

black. 4th tarsal segment of hind leg black. Brazil, Ecuador and

PS PUN a URS Saas aS ea eh be anton tly, de: migrifemur, (De SP.

4. Metapleurum and first abdominal tergite shiny. Ovipositor 1.1] as long as

hind tibia. Scutellum wholly brownish-red. Malar space 0.4 the basal

width of mandible. Tip of abdomen black or with a fine yellow line.

Sternites 5 and 6 with black marks. Bolivia. ... 6. terani Porter

S. Gupta: Prosthoporus (Ichneumonidae) QA47

Metapleurum and first tergite dull granulose. Ovipositor 1.3x as long as

hind tibia. Scutellum wholly yellow or only its sides yellow, not brown-

ish-red. Malar space 0.25x the basal width of mandible. Tip of ab-

domen ‘marked: with white.) Sternites 5-6 white ... 66055. ; Bb. 5

5. Nervulus basad of basal vein by 0.25 its length. Face with a black mark

just above the clypeal groove. Frons, vertex, temple, scape and basal

flagellar segments black. Thorax and propodeum without any marks.

Scutellum laterally yellow. Panama ....... 3. panamensis, Nn. sp.

Nervulus opposite basal vein. Face yellow, without any black marks. Frons,

vertex, temple, scape, and basal flagellar segments brownish-red.

Scutellum, a mark below wings and a semicircular mark on the apical

slope of propodeum, yellow. Brazil... ..... 4. merjoriae, Ni. Sp.

1. Prosthoporus townesi, n. sp. (Fig. 4)

Female: Face granulose with a few fine punctures just above the clypeus,

as wide as high. Clypeus granulose, its apical margin concave. Malar space

granulose, 0.6x the basal width of mandible. Temple mat. Frons granulose.

Vertex shiny mat, with fine punctures. Pronotum shiny mat, its groove finely

striate. Mesoscutum shiny mat. Prepectal carina 0.6x its length. Scutellum

shiny. Mesopleurum and metapleurum granulose with fine trans-striae. Nervulus

basad of basal vein. Second recurrent vein meeting areolet at its apical 0.4.

Abdomen mat, second tergite granulose with fine punctures sub-apically. Length

of ovipositor equal to the length of hind tibia.

Reddish-brown. Head reddish-brown, flagellar segments 1-4 black, 5-10

white, and rest black. Base of mandible, base of clypeus, and orbital mark

on frons and face, whitish-yellow. Pronotal collar, tegula, subtegular ridge,

scutellum, a mark on hind corner of mesopleurum and a transverse mark on

the apical area of propodeum (extending to 0.5 the width of metapleuron),

whitish-yellow. Wings hyaline with subapical and median brown bands. Fore

leg yellowish-brown, its apital tarsal segments darker. Middle leg reddish-

brown, tibia black, its basal end whitish, basal tarsal segment white, rest black.

Hind coxa reddish-brown, trochanters and femur brown, tibia black, its basal

end whitish, basal tarsal segment white, rest black. Abdominal tergites 1-3

and 7 apically white. The rest apicolaterally white. Sternites 2-4 brown, last

broadly white.

Male: Unknown.

Length: ll mm. Fore wing 8 mm. Ovipositor 4 mm.

Holotype: Female, Brazil: Represa Rio Grande, Guanabara. No.v 1966,

M. Alvarenga (Townes). Paratype: 1F, same locality as the holotype, Aug.

1969, M. Alvarenga (Townes).

Distribution: Brazil.

2. Prosthoporus striatifrons, n. sp.

Female: Face granulose with fine punctures, wider than high. Clypeus

granulose basally and shiny apically, its margin concave. Malar space granu-

lose, 0.5x the basal width of mandible. Temple mat with scattered fine punc-

tures. Frons shiny with fine trans-striae. Vertex shiny, mat, with fine

punctures. Pronotum shiny, mat, with fine striae in its groove. Mesoscutum

248 Contrib. Amer. Ent. Inst., vol. 20, 1983

dull mat. Prepectal carina 0.6x the height of mesopleurum. Scutellum shiny.

Mesopleurum and metapleurum granulose with fine transverse striae, sometimes

striae a little irregular. Nervulus basad of basal vein. Second recurrent vein

meeting areolet at its middle. Abdomen shiny, mat. Second abdominal ter-

gite granulose. Length of ovipositor equal to the length of hind tibia.

Reddish-brown. Color essentially same as of P. townesi, but without brown

bands on the fore wing. Tibia of middle leg, trochanters and femur of hind

leg, and sternites 2-4 lighter in color.

Male: Essentially similar to the female but frons dull mat, without any

striations, pronotal groove striate to almost smooth, mesopleurumand meta-

pleurum granulose to finely granuloso-striate, and nervulus opposite basal vein.

The color of male is rather different from that of the female. Black,

face ( except clypeal groove, a mark in the center of face, tip of mandibles,

frons and vertex), yellow. Pronotal collar, a line on upper margin of pronotum,

tegular, subtegular ridge, scutellum, a small mark on speculum, a mark on low-

er corner of mesopleurum a mark below wing, a mark on apical area of meta-

pleurum and a mark on apical side of propodeum towards metapleuron, yellow.

Wings hyaline. In fore and middle legs coxa, first trochanter and tarsal seg-

ments yellow, second trochanter femur and tibia brownish-yellow. Hind coxa

black apically with yellow mark, trochanters black, hind femur, tibia and basal

tarsal segment brown except tibia basally yellowish, 2-4 tarsal segments yellow.

Abdominal tergites 1-5 with yellow apical band.

Length: F, 7-12 mm. Fore wing 5~8.5 mm. Ovipositor 4.5 mm.

Holotype: Female, Brazil: Pedra Azul, M. G., Nov. 1970, F. M. Oliveira

(Townes). Paratypes: 5 M, Brazil: same locality as the holotype, Nov.

1972, Alvarenga and Seabra (Townes).

Distribution: Brazil.

3. Prosthoporus panamensis, n. sp. (Fig. 7)

Female: Face mat with a few fine punctures, slightly wider than high.

Clypeus shiny mat with a few punctures basally, its apical margin concave.

Malar space granulose, 0.25 the basal width of mandible. Mandible stout,

punctate at base. Temple shiny mat with a few fine punctures at its upper

corner. Mesoscutum and scutellum shiny. Prepectal carina 0.6x its length.

Mesopleurum and metapleurum granulose with fine striations in between. Ner-

vulus basad of basal vein. Second recurrent vein meeting areolet at its apical

0.4. Abdomen mat, second tergite granulose with a few fine punctures. Length

of ovipositor much longer than the length of hind tibia (1.3).

Thorax brownish-red, abdomen black with yellow stripes. Head black with

yellow marks. Flagellar segments 4-9 white. Face clypeus, base of mandible

and orbital ring white. Clypeal fovea, a mark on the center of face above

the clypeus, black. Apical slope ot clypeus with a brown mark. Thorax

brownish-red only sides of scutellum yellowish. Wings hyaline. Fore and mid-~

dle legs of same color as the thorax, tarsal segments only slightly darker.

Hind leg brownish-red, femur apically brownish black. Hind tibia black, basally

brownish-red. Tarsal segments 1-4 white, basal segment basally black. All

abdominal tergites black with apical white band, basal 0.5 of first tergite

brownish-red. Tergite 8 with an elongated white mark. All abdominal sternites

apically white.

S. Gupta: Prosthoporus (Ichneumonidae) 249

Male: Rather similar to the female, but smoother mesopleurum and meta-

pleurum without fine striations. Nervulus opposite basal vein.

Black with yellow marks. Head yellow except apex of mandibular teeth,

frons medially and vertex. Pronotal collar, upper margin of pronotum, tegula,

subtegular ridge, mark on mesothorax, scutellum, speculum, lower portion of

mesopleurum,mesosternum, a large mark below wing, metapleurum,and a small

mark on apical part of propodeum towards metapleurum, yellowish-white.

Wing hyaline. Legs brownish-yellow. Fore coxa and trochanters white. Mid-

dle tarsal segments brown. Hind trochanters, tibia and basal tarsal segments

brown. Abdomen black with white apical bands.

Length: F, 8-l4 mm. Fore wing 5.0-9.5 mm. Ovipositor 7 mm.

Holotype: Female, Panama: Darien, 1967, C. A. Triplehorn (Townes).

Paratype: M Panama: Sta. Clara Chiriqui, M, May 31, 1977. H. A. Howden

(lownes). 1 M, same data as the holotype (Townes).

Distribution: Panama.

A male from Brazil in D.P.I. Coll. probably belongs here.

4. Prosthoporus marjoriae, n. sp. (Fig. 6)

Female: Face mat, slightly wider than high. Clypeus shiny mat, its apical

margin slightly concave. Malar space mat, 0.25x the basal width of mandible.

Mandible with fine punctures. Temple shiny. Frons smooth and shiny. Ver-

tex smooth and shiny, not punctate. Pronotum shiny mat, striations absent

in the groove. Mesoscutum shiny mat. Prepectal carina 0.5x its length. Scu-

tellum smooth and shiny. Mesopleurumand metapleurum granulose, with only

a tew fine striations along metapleural fold. Nervulus almost opposite basal

vein. Second recurrent vein meeting areolet at its middle. Abdomen mat.

Second abdominal tergite granulose without punctures. Length of ovipositor

1.3x as long as the length of hind tibia.

Body brownish-red, abdomen black with yellow bands. Scape and first

flagellar segment brownish-red. Flagellar segments 4-8 yellow, rest brownish-

black. Head brownish-red. Face, clypeus, base of mandible and orbital ring

yellow. Tegula, subtegular ridge, scutellum and an inverted v-shaped mark on

the apical slope of propodeum, yellow. Wings hyaline. Fore and middle legs

yellowish-brown, only tarsal segments a little darker. Hind coxa and femur

brownish-red, trochanters brown. Hind tibia black its basal 0.3 yellowish

brown. Hind tarsal segments white. Tergite 2-8 black with apical yellow band.

First tergite brownish-red with apical yellow band. Tergite 8 with elongated

yellow mark. Sternites 5-6 yellow.

Male: Unknown.

Length: 14 mm. Fore wing 10 mm. Ovipositor 7 mm.

Holotype: Female, Brazil: Caruaru, 900 m., April 1972, M. Alvarenga

(Townes).

Distribution: Brazil.

250 Contrib. Amer. Ent. Inst., vol. 20, 1983

5. Prosthoporus nigrifemur, n. sp. (Fig.5)

Female: Face granuliose with fine punctures laterally. Face as wide as

high or slightly wider. Clypeus dull granulose. Malar space granulose, 0.25

or less than the basal width of mandible. Mandible mat, without fine punc-

tures. Mandibular teeth almost equal to lower tooth shorter. Temple frons

and vertex dull mat without fine punctures. Pronotum dull mat without fine

Striations. Mesoscutum mat. Scutellum shiny mat with a few superficial

punctures. Prepectal carina 0.6x its length. Mesopleurumand metapleur um

granulose. Propodeum transversely striate but sometimes striations weaker

medially. Nervulus basad of basal vein, second recurrent vein meeting

areolet at its basal 0.4 to apical 0.4. Abdominal tergites mat. Ovipositor

1.3x as long as the length of hind tibia.

Body brownish red, abdomen black with white stripes. Scape and flagel-

lar segments black except 5-7 white. Head black, except face laterally,

clypeus, base of mandible and a line on the frons along the eye margin,

yellow. Thorax without mark. Wings hyaline. Fore and middle legs, body

colored except tarsal segments a little darker. Hind coxa and trochanters

brownish red, femur black. Hind tibia black, its basal 0.3 yellowish-brown. _

Tarsal segments |-3 white, basitarsus basally black. Tarsal segment 4th and

claws black. Abdominal tergites 1-7 with white apical bands sometimes 8th

also. Tergite one basally brownish-red, medially black, sometimes. Abdomin-

al sternite laterally black.

Male: Similar to female. Black face, clypeus mandibles except apex, malar

space, and a line on frons aleng eye margin, whitish-yellow. Pronotal collar,

tegula, subtegular ridge, scutellum, speculum, a mark on hind corner of meso-

pleurum,mesosternum and upper half of metapleurum whitish-yellow. Wings

hyaline. Fore and middle, coxa and trochanters white; femur and tibia

yellowish-brown ventrally and brown dorsally; trochanters brownish-black.

Femur yellowish-brown, apically brown. Tibia and tarsus brown, tibia

yellowish-brown basally. Abdomen brownish-black, tergite 1-6 apically with

white stripe.

Variations: This species shows variations. The size of mandibular teeth,

which are equal or the lower tooth distinctly shorter. The junction of second

recurrent vein with the areolet also varies in position. The propodeal stria-

tions are moderately strong to weak.

In females, the color of first abdominal tergite is almost black to only

sub apically black. One species from Mangaratiba, the mesoscutum is black.

Similarly in males, the yellow mark on metapleurum may be large or small

and the color of the hind tarsal segments is yellow to black.

Length: F, 8-l0 mm. Fore wing 6-8 mm. Ovipositor 4.0 -4.5 mm.

Holotype: Female, Brazil: Represa Rio Grande, Guanabara, Dec. 1967,

M. Alvarenga, (Townes). Paratypes : 2 F and 16 M, Brazil: same locality

as holotype 1M Dec. 1967; 1 M, Aug. 1969; 3 M, Sept. 1969, all collected by

M. Alvarenga. Rio de Janeiro, | M, March 7, 1966, H. and M. Townes.

Teodoro Sampaio, | M, Nov. 1977; Bahia, Encruzilhada, 980 m, 2 M No. 1974,

M. Alvarenga, Mangaratiba, Muriqui R. D. J., 1 F, July 1969; Silva Jardin,

R...de J.,.1.F, 2.M,:Aug..1974.,...F.. Ms. Oliveira... Amazonas, 71°38). W, 4°..33'S,

S. Gupta: Prosthoporus (Ichneumonidae) 251

3 M, Sept. 1979, Alvarenga. Peru: Quincemil, 750 m, nr. Marcapata, 1 M,

Nov. 1962, Luis Pena. Ecuador: Coca, 1 M, May 1965, Luis Pena (Townes).

Distribution: Brazil, Peru and Ecuador.

SG a EDITS

Figs. 1-3. Prosthoporus terani Porter. coe

1, Lateral view showing habitus and color pattern. 2, areolet. 3, ovipositor tip.

(After Porter)

6. Prosthoporus terani Porter

Prosthoporus terani Porter, 1976. Psyche, 83: 274. F. des., fig.

Type: F, Bolivia: Santa Cruz (DPI, Gainesville). Type examined.

Female: Face mat, slightly wider than high. Clypeus shiny mat, its apical

margin slightly concave. Malar space mat, 0.4 the basal width of mandible.

Mandible with fine punctures. Temple shiny mat. Frons and vertex shiny, with

some weak shagreening. Pronotum shiny mat, without striations. Scutellum

smooth and shiny. Mesopleuramshiny mat. Metapleurum smooth and shiny.

Nervulus basad of basal vein. Second recurrent vein meeting areolet at its

basal 0.4. All abdominal tergites shiny mat. Length of ovipositor 1.1 as long

as the length of hind tibia.

Body brownish-red, abdomen black with white bands. Scape and flagellar

segment 1-3 blackish-brown, 4-9 white, and rest brown. Face clypeus, man-

dibles and orbital ring, yellow except a mark in the center of face above

clypeus and clypeal grooves yellowish-brown. Frons black. Vertex black, medial-

ly brownish-red. Wings hyaline. Fore and middle legs body colored only,

claws a little darker. Hind coxa, trochanters, femur of body color, apical 0.3

of tibia black, tarsus white. All abdominal tergites black with white apical

bands except first tergite which is basally and medially brownish-red. Ter-

gite 8 with a fine white line at its apical margin. Sternites 5-6 with black

marks.

252 ; Contrib. Amer. Ent. Inst., vol. 20, 1983

P. townesi

P. striatifrons

- panamensis

P. marjoriae

P. nigrifemur

Oe ® wm P

P. terani

Distribution of species of Prosthoporus

Male: Unknown.

Length: 12 mm. Fore wing 9 mm. Ovipositor 5 mm.

Specimen examined: Bolivia: Santa Cruz: General Saavedra Fstacion

Experimental Agricola, 1 F (type), Nov. 1973, in Malaise trap, C. C. Porter

(Gainesville).

Distribution: Bolivia.

REFERENCES

Porter, C. C. 1976. Prosthoporus, a New Bolivian Genus of the subtribe

Lymeonina. (Hymenoptera, Ichneumonidae). Psyche 83: 271-276.

Townes, H. 1970. The genera of Ichneumonidae. Part 2. Mem. Amer. Ent.

Inst. 12: 1-537. (Gabuniini, pp. 325-344).

S. Gupta: Prosthoporus (Ichneumon

idae)

253

. marjoriae.

5, Head and thorax of

. panamensis.

6, Side view of thorax of P

lor pattern.

ing co

femur show

7, Side view of thorax of P

Figs. 4-7.. 4, Fore wing of Prosthoporus townesi.

- nigri

A NEW SPECIES OF BIOBLAPSIS(ICHNEUMONIDAE: DIPLAZONTINAE)

FROM NORTH AMERICA

Michael G. Fitton

Department of Entomology, British Museum (Natural History)

Cromwell Road, London SW7 5BD, England

The genus Bioblapsis has been known only from a single, distinctive

European species. The discovery of a second species in North America neces-

Sitates a reappraisal and redefinition of the genus.

Genus BIOBLAPSIS Foerster

This new diagnosis is modified from that of Townes (1971: 229), which

also was not entirely accurate with regard to the then known species (Fitton

and Rotheray, 1982: 316). Townes (1971: 230, 358) gives a full generic synony-

my and a habitus figure.

Fore wing 3.9 to 6.6 mm long. Face 2.1 to 2.6 as wide as high, polished

or coriaceous, medially protuberant, the median protuberance not delimited

by grooves. Subocular sulcus present or absent. Clypeus 2.1 to 2.3 as wide

as high, separated from the face by a groove, polished or sub-polished, almost

flat in profile, slightly concave apico-laterally, with the edge thin and slightly

reflexed, its apical margin with a weak median notch. Antenna 0.63 to 0.92

as long as fore wing. Flagellum with 17 to 19 segments; second segment 1.7

to 3.1 as long as broad; placoid sensillae small and numerous; and with erect

bristles distinct from the clothing hairs. Male flagellum without tyloids.

Thorax polished, with scattered punctures. Notaulus indicated by a faint im-

pression. Epicnemial carina complete but sometimes obsolete ventrally.

Sternaulus absent. Prescutellar groove with longitudinal carinae, including ©

about six strong ones. Propodeum with strong carinae, its spiracle enlarged

and with an elevated rim. Petiolar face of propodeum steeply sloping. Fore

wing with 3r-m absent and cu-a slightly distal to M & Rs. Hind wing with

cu-a, shorter than abscissa of Cul between M + Cul and cu-a, and | to 3

costal hamuli. Tibiae with numerous strong hairs. Gaster more or less com-

pressed from segment 3 to apex in female, subcylindrical in male, polished.

Posterior edges of tergites 3 to 8 (F) or 3 to 7 (M) extending further back-

wards laterally than medially, especially in female. First tergite 1.18 to 1.60

as long as wide, with dorsolateral and ventrolateral carinae strong and latero-

median carinae well developed to virtually absent. Spiracle of first tergite

enlarged, but slightly smaller than that of propodeum. Second tergite 0.60

to 1.10 as long as wide, with a fold separating the laterotergite, and with the

spiracle on the main part of the tergite. Third tergite with a fold separat-

ing the laterotergite only at its extreme anterior.

1. Bioblapsis henryi, n. sp. (Fig. 1)

Female Fore wing length 3.9 mm. Face 2.1 as wide as high, coriaceous

with scattered punctures which are not easily distinguished from the general

sculpture. Median protuberance of face slightly less strong than in B. polita.

Subocular sulcus absent. Clypeus 2.3 as wide as high, sub-polished and more

weakly sculptured than face. Antenna 0.92 as long as fore wing. Flagellum

with 18 segments (the final one partially subdivided); second segment 3.1 as

long as broad; relative lengths of first, second and third segments 1:0.72:0.66;

254

Fitton: Bioblapsis (Ichneumonidae) 255

the erect bristles relatively less numerous and conspicuous than in B. polita

F. Notaulus indicated by an extremely weak impression. Epicnemial carina

complete. The longitudinal carinae in the prescutellar groove less strong

than in B. polita. Propodeal spiracle enlarged, but slightly less so than in

B. polita. Fore wing with 2r-m subequal in length to the abscissa of M

between 2r-m and m-cu. Hind wing with 2 or 3 costal hamuli (the holotype

has 2 on the left and 3 on the right). Gaster relatively short, moderately

compressed from segment 3 to apex; polished, from tergite 3 with fine, regular

alutaceous sculpture, with relatively few fine, short hairs. Posterior edges of

tergites 3 to 8 extending further backwards laterally than medially. Tergite

| 1.18 as long as wide, with the lateromedian carinae virtually absent. Ter-

gite | and the base of tergite 2 granulate. Tergite 2 0.60 as long as wide.

Apex of ovipositor sheath acute.

Blackish. Antennal flagellum brown. Whitish: spot on center of face,

clypeus, mandibles except teeth, palpi, tegula and mesepimeron dorsally.

Gaster with tergite 1 postero-medially and the posterior half of tergite 2 red-

dish. Remainder of gaster dark brown, not as black as head and thorax. Legs

reddish. Extreme base and apical third of hind tibia and most of hind tarsus

infuscate. Fore coxa and fore, mid and hind trochanters and trochantelli

whitish.

Male: Unknown.

Holotype; Female, USA: Michigan, Ann Arbor, 17. ix ~ 2. x. 1975, M. G.

Fitton [British Museum (Natural History) ].

The description of a new species from a single specimen needs some justi-

fication. In this case the most important reasons are as follows. The genus

includes only one other species, which has a relatively limited distribution and

a distinctive morphology. The new species is from a different zoogeographical

region and superficially it does not closely resemble the other species. No

material of the new species was found in collections in the course of a study

leading to a monographic revision of the subfamily Diplazontinae in North

America (Dasch 1964). The holotype was collected in a Malaise trap in the

immediate vicinity of the American Entomological Institute at Ann Arbor and

despite intensive collecting in this area over many years by Henry and

Marjorie Townes no other specimens have turned up. There is, therefore, a

need to alert others to the existence of the species if further specimens, in-

cluding males, are to be found.

To facilitate comparison a description of the European species, B. polita,

is given below. A synonymy is also included because this species is still some-

times referred to by the invalid name flavipes (for example, by Thirion 1981:

90).

2. Bioblapsis polita (Vollenhoven) (Fig. 2)

Bassus flavipes Holmgren, 1858 (1856). K. Svenska. Vetensk. Akad. Handl.

(N.F.) 1: 356. Type(s) M, Sweden: Smaland, Anneberg (? Stockholm) [not

examined]. Name preoccupied in Bassus by Lucas, 1849.

Trichomastix polita Vollenhoven, 1878. Tijdschr. Ent. 21: 161. Holotype

F, Netherlands: Scheveningen (? lost) [not examined].

256 Contrib. Amer. Ent. Inst., vol. 20, 1983

Figs. 1-2. Gaster of female, left lateral view: 1, Bioblapsis

henryi. 2, B. polita ( not to the same scale )

Fitton: Bioblapsis (Ichneumonidae) 257

Bassus tibialis Bridgman, 1883. Trans. Ent. Soc. Lond. 1883: 170.

Syntype F, Great Britain (Norwich) [examined]. |

Trichomastix pallipes (Holmgren): Thomson, 1890. Opusc. Ent. 14:

1473. Incorrect subsequent spelling of flavipes Holmgren.

Female: Fore wing length 5.3 to 6.6 mm. Face 2.3 to 2.6 as wide as high,

polished with scattered punctures and sometimes with very faint traces of

sculpture. Subocular sulcus present and relatively strong. Clypeus 2.1 to 2.3

as wide as high, polished. Antenna 0.63 to 0.66 as long as fore wing. Flagel-

lum with 17 to 18 segments; second segment 1.7 to 1.9 as long as broad; rela-

tive lengths of first, second and third segments about 1:0.62:0.55; the erect

bristles long and conspicuous. Notaulus indicated by a faint impression.

Epicnemial carina complete but sometimes obsolete ventrally. Propodeal

spiracle considerably enlarged, with its elevated rim pale. Fore wing with

2r-m very short, less than half length of abscissa of M between 2r-m and

m-cu. Hind wing with 2 (or sometimes 1) costal hamuli. Gaster relatively

long, strongly compressed from segment 3 to apex; polished, from tergite 3

with extremely fine, regular alutaceous scupture, especially laterally, with

relatively few fine, short hairs. Posterior edges of tergites 3 to 8 extending

much further backwards laterally than medially. Tergite | 1.35 to 1.60 as long

as wide, with the lateromedian carinae weak. Tergite | and the base of ter-

gite 2 longitudinally striate. Tergite 2 0.96 to 1.10 as long as wide. Apex

of ovipositor sheath rounded.

Blackish. Whitish: clypeus (especially medially), base of mandibles, palpi,

humeral bar, tegula, scutellum laterally and mesepimeron dorsally. Legs red-

dish, hind tibiae and tarsus slightly infuscate and with bases of all tibiae and

hind basitarsus whitish.

Male: As female except: Fore wing length 6.0 to 6.2 mm. Face 2.2 as

wide as high. Clypeus 2.3 as wide as high. Antenna 0.85 as long as fore wing.

Flagellum with 19 segments; without tyloids; second segment 2.8 as long as

broad; relative lengths of first, second and third segments about | : 0.72 : 0.64;

the erect bristles shorter and less conspicuous than in female. Propodeal

spiracle slightly less enlarged than in female. Gaster relatively long, sub-

cylindrical in form from segment 3, tapering slightly to apex; polished, finely

and fairly evenly punctate with longer and more abundant fine hairs than fe-

male. Posterior edges of tergites 3 to 7 extending only slightly further back-

wards laterally than medially. Tergite | 1.47 to 1.50 as long as wide, with

-the lateromedian carinae stronger than in female. Tergite | rugulostriate and

the base of tergite 2 striate. Tergite 2 1.00 as long as wide.

Blackish. Whitish: spot on center of face, facial orbits, clypeus, mandi-

bles except teeth, palpi, humeral bar, tegula, subalar prominence, scutellum

laterally and mesepimeron. Legs reddish, fore coxa and trochanter and hind

tibia and tarsus paler and with bases of all tibiae and hind basitarsus whitish.

Material examined: 8 F, 2 M, all from the British Isles. Dates of cap-

ture range from mid-June to October.

258 Contrib. Amer. Ent. Inst., vol. 20, 1983

DISCUSSION

B. henryi can be distinguished easily from B. polita by its overall

superficial appearance and there is little point in giving a key to separate them.

Apart from a few characters, such as the presence or absence of a subocular

sulcus, the differences between the species are in the degree of development

of characters. Many of these characters, such as the enlargement of the pro-

podeal spiracle, have been used, particularly in keys, to distinguish the genus

and they are less well developed in the new species. 8B. henryi will not run

to Bioblapsis in Townes key to genera (1971) nor in my own (Fitton and

Rotheray 1982). No simple modification of either of these keys would enable

it to be included. Study of B. henryi has reinforced my opinion that the

present generic classification of the Diplazontinae is unsatisfactory. Recent

work has led to more satisfactory groupings of species within genera but the

relationships between these species groups and the level at which genera should

be recognized are in need of much further study. It would be premature to

make any more comments here.

REFERENCES

Dasch, C. E. 1964. Ichneumon-flies of America North of Mexico: 5. Sub-

family Diplazontinae. Mem. Amer. Ent. Inst. 3.

Fitton, M. G. and Rotheray, G. E. 1982. A key to the European genera of

diplazontine ichneumon-flies, with notes on the British fauna. Syst. Ent.

7: 311-320.

Thirion, C. 1981. Ichneumonidae (Hymenoptera) nouveaux pour la faune Belge.

Ann. Soc. Roy. Zool. Belg. 112: 89-92.

Townes, H. 1971. The genera of the Ichneumonidae, part 4. Mem. Amer.

Ent. Inst. 17.

ICHNEUMONOIDEA (HYMENOPTERA) FROM THE

LOWER CRETACEOUS OF MONGOLIA

Alexander P. Rasnitsyn

Paleontological Institute, Academy of Sciences of the USSR

Moscow 117321, USSR

The hymenopterous superfamily Ichneumonoidea is known paleontologically

from the Neocomian, a lower part of the Lower Cretaceous, and all the olde-

est representatives described belong to the Ichneumonidae (Townes 1973;

Rasnitsyn 1975). Two new fossil ichneumonoids were collected by the Paleo-

entomological Party (Dr. A. G. Ponomarenko, Head) of the Joint Soviet-

Mongolian Paleontological Expedition. They are probably older than those men-

tioned above. They were found at Khutel-Khara-Ula (known also as Khara-

Khutul-Ula) Mts., 70 km SW of Sain-Shand, East Gobi Dsitrict of Mongolia,

in a lens of silstone and mudstone enclosed in basalts dated radiologically as

Lower Cretaceous (Shuvalov 1979). Paleoentomological data indicate an age

older than that of all previously known Lower Cretaceous insect assemblages,

because the majority of the insects in Khutel-Khara is typical for the Neo-

comian fauna as it is described by Zherichin (1978) and Rohdendorf and

Rasnitsyn (1980), while a few groups (e.g., limnic stoneflies and hemipterous

family Progonocimicidae) are absent or rare since Jurassic. The Ichneumonoid-

ea from Khutel-Khara are described below. The types are deposited in the

Paleontological Institute, Academy of Sciences of the USSR, Moscow.

PRAEICHNEUMONIDAE, new family

Figure 1

Fore wing with costal space narrow but distinct, RS+M, 2 ~3r-m and

2m-cu equally well developed, cells 2-3rm large, not modified. Hind wing

with cell r closed, short, r-m short, distant from both RS and M bases.

Mesoscutum with median suture much reduced while parapsides (notaulices)

strong. Metasoma with tergum I broad, terga I -III overlapping.

Type and the only genus Praeichneumon, n. gen.

PRAEICHNEUMON, new genus

Habitus stout. Size small. Head transverse, eyes moderately large,

temples weak. Pronotum short centrally. Mesonotum transverse with

parapsides complete, deep, narrow cephalad, broad caudad, separated postero-

medially. Median line very thin, reaching neither fore mesonotal margin nor

transverse suture. Transverse suture thin, complete. Prescutellar fovea in

form of an inverted V. Scutellum transverse with sides subparallel. Pro-

prodeum possibly areolated in the typical Ichneumonid type. Its hind slope

obscured in the fossil. Wings as figured. Legs short. Metasomal tergite |

transverse, scarcely narrower than the following, thickened and rugose except

laterally and apically. Terites If-VI transversely depressed basally or (at

least tergites II-III) subbasally, depression being strongest in tergite II and

weakest in VI. Ovipositor sheath short, narrowed apically, convex ventrally,

scarcely concave dorsally, rounded apically. Ovipositor straight (apex missing

in fossil).

259

: >

Imm 2 we

Eobracon inopinatus.

261

FIGS. 3 - 4. Praeichneumon townesi, photos

3, in polarized light. 4, in normal oblique light

262 Contrib. Amer. Ent. Inst., vol. 20, 1983

FIG. 5. Eebracon inopinatus

Photograph in polarized light.

Rasnitsyn: Fossil Ichneumonoidea 263

The type and only species, P. townesi, n. sp.

The generic name is from Ichneumon. Gender masculine.

Praeichneumon townesi, n. sp. (Figs. 1, 3, 4)

Female: Macrosculpture not mentioned above absent. Less coarse sculp-

ture discernible only in transverse subbasal depression on tergite II in the

form of fine longitudinal striation.. Head and thorax dark. Hind femur except

apex, metasomal tergite I and apices of tergites II-V less dark. Middle femur

except apex, rest of tergites II-V and tergite VI still less dark. Veins,

pterostigma, middle and hind tibiae and femoral apices, tergite VII and ovi-

positor sheath rather light. Wing membrane and tergite VIII light.

Length: Body 5.0 mm. Thorax 1.3 mm. Hind femur 0.7 mm. Fore wing

3.2 mm. Ovipositor sheath 0.25 mm.

Holotype: Female, no. 3965/416, Khutel-Khara-Ula Mts., SE Mongolia,

locality 300-1, Lower Cretaceous (Moscow).

The species is named in honor of Dr. Henry Townes.

Position and affinities: Praeichneumon is included in Ichneumonoidea

because it has autapomorphies of the superfamily (costal space reduced) and

of the infraorder Ichneumonomorpha (median mesonotal suture reduced and

first abscissa of M straight in hind wing) and lacks apomorphies of other

ichneumonomorph superfamilies (for details see Rasnitsyn 1980). A new

family is erected for this fossil because of the presence of important aut-

plesiomorphies (3rm cell closed in fore wing, r cell closed and short and short

in hind wing) and the absence of the apomorphies of other ichneumonoid

families. The family is treated as lacking apomorphies, supposing those seen

in the fossil (short oviposition, narrow temples, fore wing without interanal

cross-veins, hind wing with cu-a long, sinuate) to be features of the genus

and not the family. Thus Praeichneumonidae is a paraphyletic unit ancestral

to all other Ichneumonoidea (I would not consider paraphyletic taxa as un-

natural ones, see for details Rasnitsyn, 1980: 6-7).

BRACONIDAE

EOBRACON, new genus

Figure 2

Habitus neither stout nor slender. Size small. Head of moderate size.

Eye large, reaching mandibular base. Temple narrow. Mandible not protrud-

ing indicating absence of clypeal fovea. Palpi long. Antenna filiform with

about 16 joints, scape a little longer than pedicel, both of subequal width and

combining of subequal length to first flagellomere. Thorax short and high.

Pronotum with vertical depression subparallel to its hind margin laterally.

Mesonotum with distinct parapsids. Scutellum long. Mesopleura with pre-

pectal carina and subventral horizontal depression joining prepectal carina and

sharply margined above. Apical slope of metanotum, rather long. Propodeum

small. Legs thin, of moderate length. Fore wing with R complete (not

fractured before RS base). Pterostigma large, long, with r-rs far beyond mid-

dle. Basal vein with RS occupying 0.4 its length. Veins adjacent to

264 Contrib: Amer. Ent, Inst. ,: vol, 29, 1983

bifurcation point of RS+M much reduced. R beyond pterostigma and RS be-

yong r-rs veru sjprt/) 2~3r-m absent. M and Cu almost reaching wing mar-

gin. M bent basally indicating former position of 2r-m. 1m-cu much reduced

apically, its direction meeting M submedially between supposed RS+M apex and

former 2r-m position. Crossvein cu-a postfurcal. Brachial cell issuing Cu be-

low its midhight, not clearly closed below it. Interanal crossvein rudiments

absent. Hind wing not preserved in fossil. Metasomal terga transverse, tergites

III], unlike the following, not separable when macerated and evidently not

overlapping. Tergite I with deep medial furrow forking basad. Only latero-

tergite I present. Sternite I not seen in the fossil, sternal halves II - VI

separated (not completely so in sternite VI), small III -IV largest, II least, VI

longest), subquadrate to subtriangular, with medioapical and/or laterobasal

angles most prominent. Ovipositor short, scarcely upcurved, sheath rather wide,

rounded apically.

Type and the only species, £. inopinatus, n. sp.

The genus name is from EWS (dawn), plus Bracon.

Eobracon inopinatus, n. sp. (Figs. 2, 5)

Female: Antenna distinctly widened subapically. Scape subconical. Pedi-

cel subquadrate. Flagellomeres elongate (basal one with length ca. 4 times

apical width, penultimate hardly 1.5 times width), smoothly jointed basally and

centrally, weakly moniliform apically. Sculpture weak or absent (not discern-

able in fossil). Head, antenna, thorax and coxae dark, legs and metasomal

sclerites less dark, veins and pterostigma rather light.

Length: Body as preserved (with metasoma swollen due to maceration)

2.8 mm. (In lifetime probably 2.3-2.4 mm). Antenna 1.6 mm. Thorax 0.75

mm. Hind trochanter and femur combined 0.62 mm. Fore wing 1.8 mm.

Ovipositor 0.55 mm.

Holotype: Female, no. 3965/17, Khutel-Khara-Ula, SE Mongolia, locality

_300/1, Lower Cretaceous (Moscow).

The species name means unexpected.

Position and affinities: Eobracon is included in the Braconidae because

of the presence of the typical Braconid synapomorphies [Tergites II -III hinged

to each other and not telescopic (overlapping), cross-vein in 2m-cu absent in

fore wing] while the apomorphies of other ichneumonoids are absent. The

affinities of the new genus within Braconidae are not clear, because many

important features are not visible in fossil, while those which are visible do

not indicate affinities with certain Braconid subgroups. Some characters seem

to be unique for Eobracon, e.g., R and RS much shortened unlike M and Cu,

R not fractured before base of RS, hind wing with reduced venation, and ter-

gite I furrowed medially. One featue (cross-vein 1m-cu far postfurcal in wing

with venation reduced and RS+M leaving basal vein subcentrally and thus sub-

horizontal) is known to me elsewhere only for atypical Neoneurini described

by Tobias and Yuldashev (1979), but otherwise those insects differ much from

Eobracon. I suppose that the latter represents an early aberrant branch of

the Braconidae. The presence of such advanced form in the lowermost Lower

Cretaceous is really unexpected, because other cretaceous Braconidae are much

younger and less advanced (Rasnitsyn 1975, 1980).

Rasnitsyn: Fossil Ichneumonoidea 265

REFERENCES

Rasnitsyn, A. P. 1975. Hymenoptera Apocrita of Mesozoic. Trans. Paleont.

Inst. Acad. Sci. USSR. Vol. 147. (In Russian).

Rasnitsyn, A. P. 1980. Origin and evolution of Hymenoptera. Trans.

Paleont. Inst. Acad. Sci. USSR. Vol. 174. (In Russian).

Rohdendorf, B. B. and Rasnitsyn, A. P. (Eds.). 1980. Historic development

of the class Insecta. Trans. Paleont. Inst. Acad. Sci. USSR. Vol. 175.

(In Russian).

Shuvalov, V. F. 1979. Basic features of the stratigraphy and paleogeography

of Jurassic and Cretaceous in Mongolia. In: Basic results of the Joint

Soviet-Mongolian Paleontological Expedition studies in 1969-1979 (Proceed-

ings of the Conference). Moscow, Acad. Sci. USSR (Paleontol. Inst.).

Pp. 16-18. (In Russian).

Townes, H. 1973. Two Ichneumonids (Hymenoptera) from the Early Creta-

ceous. Proc. Ent. Soc. Washington 75(2): 216-219.

Tobias, V. I. and Yuldashev, E. Yu. 1979. Three new morphologically peculiar

species and two new genera of Braconids-neoneurins (Hymenoptera,

Braconidae, Euphorinae, Neoneurini). Trans. Zool. Inst. Acad. Sci. USSR.

88: 95-102. (In Russian).

Zherichin, V. V. 1978. Development and changes in Cretaceous and

Caenozoic faunistic assemblages (Tracheata and Chelicerata). Trans.

Paleont. Inst. Acad. Sci. USSR. Vol. 165. (In Russian).

TERSILOCHINAE AS PARASITOIDS OF INSECT PESTS OF

WINTER RAPE (HYMENOPTERA : ICHNEUMONIDAE)

Josef Sedivy

Research Institute for Crop Production

Praha—Ruzyné, CSSR

Abstract

From 1951 to 1982, eight species of ichneumon flies of the subfamily

Tersilochinae, were discovered which parasitize the coleopterous pests of winter

rape in Czechoslovakia. Of these, only Tersilochus fulvipes Grav. showed

a significant influence on the population dynamics of its host. The parasitiza-

tion by the other ichneumon fly species was low and showed considerable

variation in different localities even within one year. In large fields of winter

rape, the parasitization of larvae of Meligethes aeneus decreased with increas-

ing distance from the edge of the field. An electrical suction trap recorded

only the dispersal of ichneumonid species dispersing above the tops on the

winter rape plants. The dispersal was most intensive during morning hours and

ceased during night hours. Yellow dishes were little attractive for most of

the species. :

The ichneumon flies of the subfamily Tersilochinae are common para-

sitoids of insect pests infesting winter rape (Brassica napus var. arvensis ).

Tersilochus spp. parasitize larvae of Ceutorhynchus spp. (Curculionidae),

Meligethes spp. (Nitidulidae) and Psylliodes chrysocephala L. (Chrysomelidae).

The members of the genus Aneuclis parasitize larvae of Ceutorhynchus.,,

Meligethes and Phyllotreta spp. Two species of Phradis parasitize larvae

of Meligathes spp. that damage winter rape or develop in wild-growing

cruciferous and other plants. In the literature, most of the ichneumon flies

have been long referred to under incorrect names. Aubert and Jourdheuil

(1958) reported several species of Tersilochinae infesting insect pests of win-

ter rape in France. Horstmann (1971, 1981) treated comprehensively the tax-

onomy of the European species of Tersilochinae and provided supplementary

data on their hosts, distribution and time of occurrence. All species para-

sitize larvae of their hosts. Only Fritzsche (1956) observed Phradis inter-

stitialis Thoms. to parasitize adults of Meligethes aeneus and M. picipes.

The present contribution supplements the knowledge of the bionomics of

ichneumon fly species commonly occurring in cultivated fields of winter rape

and evaluates their impact on the populations of their host pest species.

MATERIALS AND METHODS

During the growing seasons of 1951 to 1982, the occurrence of the

ichneumon flies of the subfamily Tersilochinae was investigated in

Czechoslovakia in winter rape fields in 1955 -1957, 1966 -1969, 1976 -1976 and

1980 -1982. The observations were largely made in the potato production areas

specialized for growing winter rape. In them, the winter rape cultivation was

about 8~-12% of the total field crops. Only the observation area in Praha-

Ruzyné is situated outside this area. The Ichneumonid species composition was

evaluated from samples obtained by three times 50 strokes made by a sweeping

net in winter rape cultures at the end of the flowering period. The samplings

were taken from the western, central and southern parts of Bohemia

266

Sedivy: Tersilochinae (Ichneumonidae) 267

(Czechoslovakia). The host-parasite ratio (H:P) was evaluated from samples

of the same size taken at weekly intervals from rape stands in which the

occurrence of the ichneumon flies and their hosts was studied throughout the

growing season. Parasitization of larvae of Meligethes aeneus was determined

by the number of first instar larvae whose dark head capsule can be seen

through the translucent epidermis of the host on its dorsal or ventral side.

The differences in the degree of parasitization of larvae of M. aeneus at the

edges and in the middle of rape fields from 10 to 120 ha in area was eval-

uated in the same way. The larvae were shaken out of rape flowers at dis-

tances of 5, 10, 25, 50, 100 and 200 m from the edge of the field. For this

study, rape fields were selected which were at least approximately square in

shape. The species of the ichneumon larvae within the host larvae was not

determined. The degree of parasitization of larvae of Ceutorhynchus napi ,

C. quadridens and C. assimilis was determined by dissecting full-grown lar-

vae at the end of the flowering period of winter rape. As for c. pleuro-

stigma of the autumn strain, the dissections of the larvae were carried out

in the spring months.

The daily flight activity of the ichneumon flies was studied using a small

Johnson~-Taylor suction trap placed in small experimental plots of winter rape

in Praha-Ruzyné. The entrance opening of the suction trap was 1.2 m above

the ground. The mechanism of this type of suction trap makes it possible to

evaluate insect samples taken at intervals of 60 minutes. The occurrence of

ichneumon flies on the phenophase of winter rape was studied using six yellow

dishes placed in the same plots as in the case of the suction trap. The posi-

tion of the dishes varied in the course of the growing season: they were

gradually lifted so that their bottom was on the level with the tops of the

rape plants.

RESULTS

Eight species of ichneumon flies are recorded here which parasitize win-

ter rape pests of the order Coleoptera in the territory of Czechoslovakia. The

periods of their occurrence, including findings outside winter rape stands, are

given in Table I.

Depending on their occurrence in winter rape stands, those ichneumon

fly species that are on the wing in March and April are the most decimated

by insecticide treatment of the rape stands which is applied at that time.

-The stands bloom in May and early June at which time no insecticides are

applied that could affect the occurrence of the parasitoids. Those species that

occur in winter rape stands in June and early July may, in some years, be

decimated by insecticide applications against overcrowded populations of the

cabbage aphis (Brevicoryne brassicae L.) infesting winter rape pods.

Tersilochus heterocerus (Thoms.), Phradis morionellus (Holmgr.)

Adults of T. heterocerus occur together with P. morionellus. Suction

trap samples, however, suggest that the period of occurrence of the latter

species in winter rape stands is shorter than that of the former Fig. as

The parasitization of larvae of M. aeneus by both species varied in successive

years as well as in different localities. In peak years of the pest and in the

subsequent years the percentage parasitization was higher (Table II). As for

268 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table I. Periods of occurrence of ichneumon flies in winter rape stands and

in natural habitats.

; Period of

Species Host

occurrence

1. Aneuclis incidens Meligethes aeneus 28.1V. — 30.1X.

(Thoms.) M. viridescens

2. Aneuclis melanarius Ceutorhynchus pleurostigma Sav. OS ZeIV iT.

(Holmgr.) C. assimiiis

Psylliodes chrysocephala

3. Phradis morionellus Meligethes aeneus 161V..— 30, Vill.

(Holmgr.) M. viridescens

| M. symphyti

4. Phradis interstitialis Meligethes difficilis ISM. <1 7 VIF

(Thoms.) M. picipes

M. viridescens

5. Tersilochus heterocerus Meligethes aeneus ZO1 V0 OVI,

(Thoms.) M. viridescens

6. Tersilochus fulvipes Ceutorhynchus napi IZAV..-— 14.VI.

(Grav.) C. punctiger

7. Tersilochus obscurator Ceutorhynchus quadridens LIV. - 30.VI1.

Aubert . Psylliodes chrysocephala

6. Tersilochus ‘tripartitus Psylliodes chrysocephala Ae Gm, BV

Brischke |

T. heterocerus, the H:P ratio showed considerable variation in successive

years: it was 18.5:1 in 1955, 8:1 in 1956, 21.5:1 in 1957, 36.8:1 in 1981 and 20:1

in 1982.

Tersilochus fulvipes (Grav.)

This species was the dominant parasitoid of larvae of c.napi. During

years of serious occurrence of C. napi the parasitization of the weevil larvae

attained 79 ~98%. Since 1977 the occurrence of C.napi has never attained

economically significant levels and the single larva found in rape stems were

mostly parasitized. The high degree of parasitization of larvae of C. napi

during years of their low numbers suggests a significant role of T. fulvipes

in the regulation of their numbers (Table III). The abundance of adult

T. fulvipes in winter rape stands was the highest in late April and early May

and was unfavorably affected by intensive chemical control of adult m.

aeneus. This species was absent from the catches in the suction trap. Oc-

casional specimens were obtained from the yellow dishes in years of higher

population densities of the host.

Sedivy: Tersilochinae (Ichneumonidae)

Fig. 1. Intensity of dispersal of ichneumon flies during 24

hours, as revealed by a Johnson-Taylor suction trap.

A , Tersilochus

obscurator. 3B, Phradis morionelius. C, Tersilochus heterocerus.

270 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table II. Parasitization of larvae of Meligethes aeneus (n = 300)

No. ichneumonid larvae per one host larva (in %)

Date Locality

0 1 2 3 4 5

5.VI.55 Praha-Ruzyné* 12.- 10.1 211 25.6 10.5 20.6

30.V.56 Praha-Ruzyn6 22.3 S.1 176. 2 5 7.6

20.V.5/ ._Praha-Ruzyne .. 10,- roe 60.2 0.8 9.3 27

12.V.66 Benesov 50.2 42.3 0.6 1.9 2.6 0.4

4.V.67 BeneSov* 20.- 29.8 51.- 0.8

22.V.69 BeneSov* 98.- 1.8 0.2

29.V.74 Benesov 92.3 1.6 6:- 0.1

19.V.75 BeneSov 94.- 6.- |

14.V.76 Tachov* 95.- 2.3 Zl

28.V.80 Novy Bor* 1h. 59.2 pes 3.3 12 0.3

30.V. Tachov 42.1 48.6 6.3 3.-

22.V.81 Praha-Ruzyné 47.- 5461 1.9

9.VI. Slapy 61.2 3911 0.7

28.V.82 Praha~Ruzyneé* 23.- 15,- 6.- Lt 41.4 Ze

*Peak years of M. aeneus, attaining economic level.

Tersilochus obscurator Aubert

The females infest 3rd instar larvae of C. quadridens. In 1955 through

1982, the parasitization determined in 18 randomly selected stands varied be-

tween 6 and 69%. During 1982 in five observation sites, the parasitization

varied within the range of 12-31%. The importance of this parasitoid for the

population dynamics of the host was small as the percentage parasitization was

low and considerably variable (Table IV).

Aneuclis incidens (Thomson)

During the study years the occurrence of this species was sporadical in

the winter rape stands. No occurrence was recorded for 1975, 1980-82. The

species is abundant on cruciferous weeds during summer.

Aneuclis melanarius (Holmgren)

The species was currently reared from the autumn strain larvae of Cc.

pleurostigma and occasionally also from those of C. assimilis. Their infes-

tation was invariably very low and did not affect the numbers of the host

species.

Sedivy: Tersilochinae (Ichneumonidae) oT"

Table III. Parasitization of larvae of Ceutorhynchus napi by Tersilochus

fulvipes

; No. larvae Neptada

Date Locality eae H:P parasitized by

T. fulvipes

5, V155 Praha-Ruzyné* 100 58-1] 88

30.V.56 Praha-Ruzyné 62 2.631 60

20.V.57 Praha-Ruzyné +2 2.6:1 10

12.V.66 Benesov 39 33

4.V.67 Benesov 18 8

22.V.69 Benesov* 100 19

29. V.74 BeneSov* 100 18.1:1 82

19.V75 Benesov_ . 32 26

14.V.76 Tachov 100 89

28.V.80 Novy Bor 23 18

30.V.80 Tachov 10 7

22.4 at Praha—Ruzyne 1 L331 1

Dav Slapy 2 2

Zi Od Praha~Ruzyneé 1 0

4.VIL. Praha—Ruzyné 1 1.3:1 1

5. VIL. Teo 2 2

*Occurrence of Ceutorhynchus napi above the economic level.

Phradis interstitialis (Thomson)

This species has been found to occur in warm years and in weedy cul-

tures of winter rape (1955, 1976, 1982). Attempts at rearing this species

from larvae of M. aeneus have been unsuccessful. The occurrence of this

species coincided mainly with the end of flowering period of winter rape.

Tersilochus tripartitus Brischke

The species is common in winter rape stands only in years of serious

occurrence of Psylliodes chrysocephala. It occurred in rather large numbers

only in 1955 and 1956. In 1956, an economically important occurrence of its

host, P. chrysocephala , was recorded. The H:P ratio was 16.2:1. In the sub-

sequent year the abundance of P.chrysocephala dropped below the economic

threshold and a H:P ratio of 4.1:1 was ascertained in the course of the season.

In the remaining years of study, T. tripartitus occurred singly and the num-

bers of its host were economically insignificant.

272 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table IV. Parasitization of larvae of Ceutorhynchus quadridens

by Tersilochusobscurator (n = 100)

ee Neale ee eo ee

S2VESS Praha-Ruzyné* 73.4:1 27

30.V.56 Praha-Ruzyné 3.534 56

20.V 37 Praha-—Ruzyné 1.6:1 82

12.V.66 BeneSov 20

4.V.67 Benesov* aaa

22.V.69 BeneSov* 69

29.V.74 BeneSov 6

19.V.75 Benesov* 48.3:1 39

14.V.76 Tachov 11

28.V.80 Novy Bor 30

s0.V: Tachov 9

22.V.81 Praha-Ruzyné 52.331 38

9.VI. Slapy 21

27.V.82 Praha-Ruzyné 19

9.Mi. Slapy 18

4.VIl. Neveklov 25

4.VII. Praha-Ruzyne 50.8:1 31

5.VII. Tachov 12

*Occurrence of Ceutorhynchus quadridens above the economic

level.

In winter rape fields larger than 50 ha the percentage of parasitized lar-

vae of Meligethes aeneus decreased with increasing distance from the edges

of the field. The degree of parasitization of the larvae was correlated with

the decrease in the average number of larvae of M. aeneus per rape plant.

The percentage of parasitized larvae was higher in a strip up to 25 m along

the edges of the stand. At a distance of 50 m from the edge and deeper in-

side the stand the parasitization of the larvae was lower and almost did not

vary. The effect of the higher parasitization at the edges of the stand was

marked in fields at least 300 m broad (Table V).

In the suction trap, Tersilochus heterocerus, T. obscurator and

Phradis morionellus were captured in May through July, 1982. T. hetero-

cerus was the most abundant: in all, 1,458 specimens were captured. The

Sedivy: Tersilochinae (Ichneumonidae) 273

flight activity of this species varied only slightly between 7 and 20 hrs. Oc-

casional dispersal was recorded during the night. The sex ratio was | male:

1.2 females. The flight intensity of Phradis morionellus was somewhat weaker

than in the foregoing species. In all, 974 specimens were captured. A rather

high flight activity was recorded between 7 and 8 hrs whereupon it decreased

gradually until 19 hrs. A single individual was found flying during night hours.

The sex ratio was | male : 4.3 females. Compared to the two foregoing

species, the flight activity of Tersilochus obscurator was the least intensive.

Increased flight intensity was recorded between 7 and 8, ll and 14, and 16 and

20 hrs. Occasional flying was recorded during night hours between 20 and 6

hrs (Fig. 2).

Table V. Percentages of parasitized larvae of Meligethes aeneus at differ-

ent distances form the edge of the field.

inh ance ihatseibelhea dae est lot cat seta eee DN eae te ee i hse es RR

Distance from edge, m

eo ameniner toh ih ii metcerie mtis Hb 9 8 4 88 Eas eT

5 10 25 50 100 200

Benesov 60 94.- 96.4 69.4 43.- $9.2 36.-

Tachov 120 10.1 7+ [3..< 0.9

Mochov 70 92.3 89.8 67.8 88.2 24.9 234%

Duba 20 23.8 Zs. 17.- 26. - 20.9 10.-

Popovice 10 Vs ee 70.- 70.8 70.8 G12 60.5

Svetice 16 85.3 84.2 82.- 80.7 76.1 73.6

The first specimens of P. morionellus were present in the suction trap

on 14 May, 1982. The numbers of this species increased rapidly and attained

a peak on 25 May. The last specimens were obtained on 4 June. In a suction

trap situated in a neighboring field of winter wheat, the last adults (both males

and females) were captured as late as 24 June. In winter rape stands,

Tersilochus heterocerus showed the longest period of occurrence. The first

adults were captured in the suction trap on 16 May; the last ones, on 6 July;

the peak numbers were recorded between 20 May and 4 June. The first adults

of Tersilochus obscurator were ascertained in the winter rape stand on 25

May, the last ones on 24 June. Their increased occurrence was recorded on

1~4 June (Fig. 2). In all the above mentioned species the most numerous oc-

currence of adults coincides with the period at which their host larvae become

full grown. Occasional adults of T. heterocerus, T. obscurator and P.

morionellus were also captured in a suction trap situated in the neighboring

winter wheat field. In that trap, most adults were captured between ll June

and 4 July; the greatest flight intensity was observed between 8 and 14 hrs.

In the yellow dishes only two ichneumon fly species parasitizing larvae

of the insect pests of winter rape were captured during April through June,

1982. Tersilochus heterocerus was numerous; increased numbers were

274 Contrib. Amer. Ent. Inst., vol. 20, 1983

log n+]

15 20 5 30] 3 5 10 15 20 25 30 5

Mai juni july

Fig. 2. Seasonal dynamics of the occurrence of ichneumon flies cap-

tured in the suction trap in 1982. A, Tersilochus obscurator. B, T.

heterocerus. C, Phradis morionellus.

J 7 ;

Sedivy: Tersilochinae (Ichneumonidae) 275

recorded on 15 -18 and 24-30 May. In all, 416 adults were captured in six

dishes, the observed sex ratio being | male:0.7 females. Phradis morionel-

lus was less numerous. Increased numbers of this species were observed on

the same days as above. A total of 51 specimens was captured in the yellow

dishes. Their sex ratio was | male:1.1 females. No other Tersilochinae

species were present in the catches in the yellow dishes.

DISCUSSION

The hitherto data on the period of occurrence and on the hosts of the

ichneumon flies parasitizing insect pests of winter rape suggest that they are

oligophaous and polyphagous species which mostly change habitats and hosts

in the course of the growing season. Exceptions are provided by Tersilochus

fulvipes and T.tripartitus whose adults occur within a rather short period

in spring at which time also the larvae of their known hosts occur. In winter

rape cultures, the abundance of species oriented to parasitize a single host

species is more numerous in the year of the mass occurrence of their host

and in the subsequent year. In years of low numbers of the hosts the occur-

rence of their parasitoids is mostly little numerous as well. The observed

species parasitizing the pests of winter rape exerted little influence on the

population dynamics of their hosts, except Tersilochus fulvipes which is

considered an effective parasitoid of Ceutorhynchus napi (Balachowsky 1963).

The percentage parasitization of larvae of this weevil was higher in years un-

favorable for the embryonic and larval development of the host. The percent-

age parasitization increased in subsequent years and the retrogradation of

C. napi deepened even under conditions favoring the development of the pest,

for example in 1982.

The parasitization of the remaining pests was considerably variable and

low. The peak numbers of the most abundant species, such as Tersilochus

heterocerus and Phradismorionellus, were observed as late as during the

period at which the winter rape ceased to flower and the host larvae had most-

ly left the plants to pupate in the soil. In such species the degree of para-

Sitization varied considerably in different localities within one year. Apparent-

ly, the parasitization of the hosts depends on the intensity of dispersal of the

ichneumon flies after hibernation. The electrical suction trap used in the

study of the dispersal of adult ichneumon flies over winter rape stands was

of only limited use, as it captured only insects flying on the level of the

entrance opening. The trap is unsuitable for recording the occurrence of

ichneumon flies moving within the stand. The suction trap mostly collected

those species which parasitize Meligethes spp. whose larvae develop in the top

parts of winter rape plants. The catches of Tersilochus obscurator, which

species parasitizes larvae of C. quadridens developing in the tissues of winter

rape stems, probably comprised adults which immigrated, or emigrated from,

the winter rape stands. The number of ichneumon flies captured in a suction

trap cannot be used to evaluate their numbers in a stand, as it only records

their flight activity above the stand. The highest flight intensity was recorded

during morning hours; the dispersal ceased during night hours.

The considerably different flight intensity of the ichneumon flies revealed

by the suction traps placed in a winter rape and a winter wheat field suggests

that the dispersal of the parasitoids was directed towards the host plants of

276 Contrib. Amer. cots, Inst.) vol 20, 1983

their host species. What factors direct the dispersal of the ichneumon flies

in search of their hosts cannot be judged from these observations. Townes

(1972) believes the preference for certain ecological niches to be of decisive

importance. From the low occurrence in, or virtual absence of certain

ichneumon fly species from, the yellow dishes one may conclude that the

yellow color of the flowering winter rape stands does not act as a visual at-

tractant that would direct the dispersal of some of the ichneumon fly species.

REFERENCES

Aubert, J., Jourdheuil, P. 1958. Nouvelle description et biologie de quelques

ichneumonides appartenant aux genres Aneuclis Foerst., Isurgus Foerst.,

et Thersilochus Holmgr. Rev. Path. Veg. et d'Ent. Agric. de France 37:

175-193.

Balachowsky, A. S. 1963. Entomologie appliquee a l'agriculture I, Coleop-

teres, Vol. 2: 1013-1021.

Fritzsche, R. 1956. Beeinflussung der Populationsdichte verschiedener

Meligethes-Arten von gleichen Wirtspflanzen durch Parasiten. Ber. uber

die Hundertsjahrfeier der DEG Berlin: 141-143.

Horstmann, K. 1971. Revision der europaischen Tersilochinae I (Hym. Ichneu-

monidae). Veroffentlichungen der Zool. Staatssammlung Muenchen 15:

47-138.

Horstmann, K. 1981. Revision der europaischen Tersilochinae II (Hym.

Ichneumonidae). Spixiana, Suppl. 4, p. 76.

Townes, H. 1972. Ichneumonidae as Biological Control Agents. Proc. Tall

Timbers Conf. on Ecol. Animal Control by Habitat Management 3: 235-

248.

SENWOT, A NEW GENUS OF ALYSIINI

(HYMENOPTERA : BRACONIDAE) FROM AFRICA

Robert A. Wharton

Department of Entomology, Texas A.&M. University

College Station, Texas 77843

Our knowledge of the interrelationships within the Alysiini is based almost

entirely on the western Palearctic fauna. Yet two-fifths of the known genera

are not found in this region. In other regions, however, most of the species

and many of the genera are still undescribed. Before significant advances can

be made in the higher classification of this tribe, much more basic descriptive

work is needed. Description of this new genus will facilitate phylogenetic dis-

cussions during revisions now in progress of Hovalysia and related genera.

SENWOT, new genus

Type~species: Senwot africanus, n. sp.

Head subcubical. Mandibles long, broadly expanded apically, with 4 teeth,

the additional tooth formed from a dorsal-apical cleft of tooth 1; teeth well-

rouned, with tooth 2 protruding only slightly beyond teeth | and 3; diagonal

ridge absent. Clypeus short, narrow, pointed, strongly protruding. First flag-

ellomere slightly shorter than second. Pronope present. Sternaulus broad, cren-

ulate, long, extending to mid coxa. Scutellum low, without posterior spine.

Metanotum with midridge, but this not elevated as a tall flange or spine. Pro-

podeal spiracle minute. Fore wing venation complete; Ist radial segment arising

before middle of narrowly elongate stigma; 2nd radial segment longer than

cuqul; brachial cell narrow but well-developed, closed at distal corner by d3;

parallel vein interstitial or nearly so; recurrent vein antefurcal to postfurcal;

nervulus antefurcal; postnervellus long, well-pigmented, strongly antefurcal;

radiella and cubitella weakly pigmented, cubitella a little longer than radiella;

Ist mediellan segment nearly as long as 2nd mediellan segment, basella short-

er than both. Tergum 2 striate; remainder of gaster unsculptured. Ovipositor

sheath moderately hairy, the hairs relatively long.

Senwot belongs to the Phaenocarpa complex of alysiine genera (Fischer

1971, Wharton 1980) because of the long second flagellomere. The linearly

elongate stigma with rl arising near the base, the absence of a diagonal ridge

on the mandible, and the antefurcal nervulus combine to readily distinguish

Senwot from all other genera in this complex.

Because Senwot is so distinctive, relationships between it and other genera

are not readily apparent. The absence of the diagonal ridge, the sculpture

of the abdomen, and the shape of the clypeus, however, suggest that its af-

finities are with Hovalysia and Hylcalosia. It must therefore be concluded

that the anterior migration of the parallel vein has occurred independently in

Senwot and the Heratemis-Phaenocarpa-Gnathopleura group of genera.

The diagonal ridge is list, reduced, or obscured in several unrelated

alysiine general. In some groups, such as Alloea and most Aspilota species,

this is correlated with a reduction in mandible size and/or disappearance of

the apical scoop. In most genera lacking a distinct diagonal ridge, the carina

forming the ventral border in the mandible is straight, indicating a simple loss

of the diagonal ridge. In Senwot, Hovalysia, Hylcalosia, and Microcrasis,

277

278 Contrib. Amer. Ent. Inst., vol. 20, 1983

however, the carina curves dorsally from the apex, then back ventrally to the

base, creating a concave ventral ridge when viewed ventral-posteriorly. Thus

suggests that the diagonal ridge has fused with the carina.

Senwot is dedicated to Dr. Henry Townes. Senwot is Townes in reverse,

and the gender is masculine.

Senwot africanus, n. sp.

Head about 1.4 times broader than long; 1.5 times broader than mesonotum.

Head in dorsal view slightly wider at eyes than at temples; eyes about twice

length of temples in lateral view. Frons, vertex, occiput, and temples with

only a few scattered hairs. Eye hairs sparse, minute, barely visible at 32X;

malar space nearly absent. Face convex; midridge broad, polished, extending

between antennal bases as a carina nearly to median ocellus, where it bifur-

cates to form a short v; surface of face transversely strigose and distinctly

punctate; face about 1.9 times wider than high. Frons with weak striae ex-

tending ventrally and laterally from midridge, otherwise smooth and polished.

Clypeus narrow, triangular, strongly protruding; epistomal sulcus deep, broad,

trough-like; face nearly 3 times wider than clypeus. Paraclypeal pits small,

distant from eye. Mandible (Fig. 2) about 1.5 times longer than apical width;

apex broadly expanded, about 1.5 times wider than base; mandible about 0.55

times length of head; original tooth | with dorsal-most is smaller; ventral—most

about equal in size to tooth 3, tooth 2 slightly larger than these, and more

distinctly triangular; upper border strongly, though asymmetrically concave, low-

er border symmetrically concave; mandible surface weakly rugose, hairy. Anten-

nae about equal in length to body plus ovipositor; 29 segmented; first flagel-

lomere about 0.8 times length of second; remaining flagellomeres relatively

long and slender, fifth flagellomere about 2.5 times longer than wide. Maxil-

lary palps 6 segmented, about 0.95 times length of thorax.

Thorax twice longer than wide. Pronotal collar with pronope as a simple,

round pit, collar otherwise smooth except for shallow, weakly crenulate trans-

verse sulcus posteriorad pronope. Mesonotum sharply declivous anteriorly;

notauli broad, foveolate, complete to broadly oval midpit. Median mesonotal

lobe hairy, with 35-40 more or less decumbent hairs covering anterior 0.66,

posterior-most hairs longer and more erect; lateral lobes with scattered, erect

hairs, mostly around border; median mesonotal lobe distinctly raised above level

of lateral lobes. Prescutellar pit with 3 evenly spaced ridges; about 2.65 times

wider than long. Scutellum sparsely hairy, flat. Metanotum with lateral fields

sculptured with longitudinal ridges. Propodeum with median longitudinal carina

over basal 1/4th, areas laterad carina largely smooth; remainder reticulate, with

reticulations more widely spaced apically. Sternaulus very broad, foveolate,

sinuate, extending posteriorly to mid coxa, and continuing uninterrupted dorsally

through subalar area; remainder of mesopleuron largely smooth, with about 10 -

12 hairs clustered ventral-posteriorly; mesopleural fovea an isolated pit. Meta-

pleuron with median unsculptured area large, with a few scattered hairs; dorsal-

posterior and ventral-posterior areas deeply excavated; anterior—ventral area

densely hairy. Tibial spurs, especially inner spur of hind tibia, hairy; hind tibia

at apex with well-defined comb medially.

Wing (Fig. 1): Stigma long, narrow, similar to members of the

Dapsilarthra apii group; roughly 7 times longer than wide; rl arising from

basal 1/3rd; r2 about twice length of rl; r3 ending at wing tip; cuqul bent,

Wharton: Senwot (Ichneumonidae) 279

roughly 0.7 times length of 12; n. rec. interstitial or very weakly antefurcal;

n. distinctly antefurcal. Hind wing with 3 hamuli; postnervellus pigmented

and sclerotized, extending basad nearly to wing margin at a 45 ~60° angle

from mediella; second mediellan segment about 1.5 times longer than Ist,

roughly 3 times longer than basella.

Abdomen: Apical width of petiole subequal to its length; apex about

twice wider than base; surface striate. Tergum 2 (Fig. 30 striate, anterior

border carinately margined except for median 1/3rd. Ovipositor sheath 0.95

times length of thorax; ovipositor roughly 1.65 times longer than thorax.

Color dark brown; mandible orange; legs, palps, scape, and pedicle yel-

low, except hind coxae brown and fore and mid coxae yellow-brown.

Length: 3.2: 3.5mm.

Holotype: Female, Nigeria: Aba, June 1967, J. Townes (Townes).

Paratype: Nigeria: Ibadan, 1 F. 16. VII. 1962, D. C. Eidt, Malaise trap

(Ottawa). Other specimens : Congo Belge (= Zaire): Yangambi, 2 M, Nov.

23, 1950, J. M. McGough (Washington).

It seems probable that the two specimens from Zaire (in USNM) are the

males of this species because of their morphological similarity. However, a

single male in the CNC from the Ibadan locality is considerably different (see

below), thus necessitating a cautionary interpretation. I have therefore labeled

the males from Zaire as this species, but not as paratypes. They differ from

the females described above as follows: Middle flagellomeres longer, more

slender; antenna 27 segmented in one specimen, broken in other; stigma long-

er, with rl arising from basal fifth; ratio of cuqul to r2=0.80; nervulus more

strongly antefurcal; scutellum not as flattened; and petiole proportions more

variable. The striae on tergum 2 are bordered posteriorly by a sulcus, but

this may be an artifact as both specimens are in poor condition. The two

males were reared during an exploration program for fruit fly parasitoids, ap-

rae from puparia in false kamani or mixed Sapotaceae (Clausen et al.

1965).

In addition to the four specimens referred to S. africanus, I have seen

a fifth specimen which apparently represents a new species. Since there is

but a single male, however, its description at this time would be premature.

It has all the features listed for the genus, but differs from S. africanus in

the shape of the head, sculptural details, and especially the venation.

ACKNOWLEDGMENTS

I am grateful to P. Kovarik for preparing Fig. 2 and to T. Friedlander

for commenting on the text. I also wish to thank the curators of the USNM,

CNC, and Townes Collection for loaning me the material used in the descrip-

tion.

REFERENCES

Clausen, C. P., Clancy, D. W. and Chock, Q. C... 1965... Biological control .of

the Oriental fruit fly (Dacus dorsalis Hendel) and other fruit flies in

Hawaii. USDA Tech. Bull. 1322.

280 Contrib. Amer. Ent. Inst., vol. 20, 1983

Fischer, M. 1971. Untersuchungen uber die Europaischen Alysiini mit

besonderer Berucksichtigung der Fauna Niederosterreichs. Polskie Pismo

Ent. 41: 19-160.

Wharton, R. A. 1977. A New World Aphaereta species (Hymenoptera:

Braconidae), with a discussion of terminology used in the tribe Alysiini.

Ann. Ent. Soc. Amer. 70: 782-803.

Wharton, R. A. 1980. Review of Nearctic Alysiini (Hymenoptera: Braconidae)

with discussion of generic relationships within the tribe. Univ. Calif.

Publ. Ent. 88: 1-112.

FIGS. 1 - 3. Senwot africanus, n. sp. Female.

1, Fore wing. 2, apical 0.6 of mandible. 3, Second abdominal tergite,

A REVISION OF THE NEW TRIBE BRULLEIINI

(HYMENOPTERA : BRACONIDAE)

C. van Achterberg

Rijksmuseum van Natuurlijke Historie

Leiden, The Netherlands

Abstract

A new tribe is erected for the genera Brulleia Szépligeti, 1904 and

Parabrulleia, n. gen. (Braconidae, Helconinae). Five new species are described;

Brulleia euphemia Turner, 1919 is synonymized with Parabrulleia shibuensis

(Matsumura 1912), n. comb.; Cenocoelius annulicornis Cameron, 191] is synony-

mized with Brulleia melanocephala Szépligeti, 1904, and Brulleia latian-

nulata (Cameron 1911) is a new combination in the genus Brulleia. The

species are keyed, described and illustrated and their phylogeny is discussed.

INTRODUCTION

The isolated genus Brulleia Szepligeti has long consisted of two valid

species: the type-species from New Guinea (B. melanocephala Szépligeti,

1904) and Doryctes shibuensis Matsumura, 1912 from Japan, China and Vietnam.

Examination of the holotype of B. melanocephala showed that the maxillary

and labial palpi both consist of two segments; however, in East Palearctic spp.

the palpi are far less reduced. Further examination of the available material

(including 5 new species) proved that the palpi do not provide reliable charac-

ters for generic division; the New Guinea species is at one end of a continuum

and the Palearctic species at the other end. Nevertheless, one species proved

to be sufficiently different to warrant the erection of a new genus. Both

Brulleia Szépligeti and Parabrulleia , n. gen., are placed in the new tribe

Brulleiini because they do not fit in the Helconini or in the Diospilini. The

Brulleiini differ from the Helconini by the absence of vein 1-SR fore wing,

the lack of a longitudinal lamella on the fronts, and the ventrally smooth hind

femur. The Diospilini differ by the antefurcal vein m-cu of fore wing, the

shorter fore tarsus and hind trochanter, the wide apex of the fore wing and

the (sub)vertical vein cu-a of hind wing. For a further analysis, see Fig. 1A.

The hosts of the Brulleiini are largely unknown; there is one record of

larvae of Cerambycidae (Coleoptera) as host of Parabrulleia shibuensis

(Matsumura) comb. nov. For technical terms used, see Van Achterberg (1979).

PHYLOGENY

The phylogenetic relationships of the species of the Brulleiini are far from

easy to understand; only one species, shibuensis, clearly deviated from the

rest. The seven known species of Brulleia occur from Japan to New Guinea

and are closely related, despite striking differences. In Table 1 the few

apomorphous character states are depicted and in Fig. 1B the most likely rela-

tionships are given. Certain character states are considered to be apomorphous

as the result of out-group comparison, mainly with the Helconini and Dios-

pilini. If the interpretation as given in Fig. 18 is correct, then Japan was

colonized from the main land, and the Philippines via Japan. Both Java and

281

1983

Contrib. Amer. Ent. inst., vol. 20,

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van Achterberg: Brulleiini (Braconidae) 285

New Guinea were colonized from the Philippines. Additionally it can be as-

sumed that intermediate species remain to be discovered in Borneo, Celebes

and the Moluccas.

BRULLEIINI, new tribe

Diagnosis: Length of fore wing 9.7 -19.0 mm, of body ll.7-24 mm;

antennal segments 33-44; antenna with no apical spine; annellus more or less

differentiated (Figs. 59, 71, 85); labrum broadly truncate (Figs. 40, 65) or

emarginate (Fig. 56); anterior tentorial pits large and deep (Figs. 6, 9); frons

slightly concave (almost flat; Fig. 52) to distinctly concave (Figs. 6, 9), with

no medio-longitudinal lamella; maxillary and labial palpi with 2-5 and 2-3

segments, respectively; face beside antennal sockets depressed (Figs. 6, 40);

occipital carina bent towards hypostomal carina, joined distinctly above base

of mandible (Fig. 20); malar suture (virtually) absent; mandible slightly twist-

ed; pronope deep (Figs. 13, 23, 33); mesoscutum setose and finely punctate;

scutellum with (rather) widely crenulated band posteriorly (Figs. 23, 34, 85);

metapleural flange medium-sized (Fig. 2) to virtually absent (Fig. 35); propodeal

spiracle round to slightly elliptical, distinctly in front of middle of propodeum

(Fig. 2); propodeum with at most a short medial carina anteriorly (Fig. 69),

with no areola or transverse carina and posterior part not differentiated from

anterior part; vein 1-SR of fore wing and vein 2A of hind wing absent (Fig.

3); veins r-m, 2A and a of fore wing present (Fig. 17); veins cu-a of both

fore and hind wing strongly inclivous (Figs. 50, 87); vein m-cu of fore wing

postfurcal (Fig. 26); marginal cell of hind wing more or less widened (Figs.

3, 26); basal and Ist subdiscal cells of fore wing (largely) glabrous (Fig. 92);

fore wing with (rather) slender apex (Figs. 3, 61, 79); vein a of fore wing is

distinctly removed from vein cu-a (figs. 3, 17); length of hind tibia 1.6 ~2.4

times hind femur; hind femur smooth ventrally; fore spur rather specialized

(Fig. 80), its length 0.2 -0.3 times fore basitarsus (Fig. ll); tarsal claws

slender, simple (Figs. 7, 34, 58); fore tarsus very long, 1.7 -2.0 times fore

tibia; inner and outer hind spurs (sub)equal, their length 0.15 -0.25 times hind

basitarsus; metasoma inserted just between hind coxae (Fig. 35) or somewhat

above coxae (Fig. 15); dorsope absent (Fig. 11); laterope large, and more or

less elliptical (Figs. 2, 35, 59); ovipositor with subapical notch (Fig. 41); length

of ovipositor sheaths 1.24 -1.9] times fore wing.

. Fs. .

Contains two genera: Parabrulleia, n. gen. and Brulleia Szepligeti, 1904.

Parabrulleia shibuensis (Matsumura), is a parasite of the larvae of

‘Cerambycidae (Coleoptera).

KEY TO SPECIES OF THE TRIBE BRULLEIINI

1. Mandibles angularly bent medially (Fig. 6); 2nd tergite (at least super-

ficially) sculptured basally (Fig. 12); Ist tergite extensively sculptured

posteriorly (Fig. 12); length of ovipositor sheath 1.60 -1.91 times fore

wing; occipital carina evenly curved dorsally (Fig. 9); length of hind

tibia 2.3 -2.4 times hind femur; clypeus (including margin) flattened

igs, Gos Aaah Lae Be i ee yuh lacte Biull erg ea ylie tan a's

ame ie Rn ae on, Mer ee vey ie 1. shibuensis (Matsumura), n. comb.

286 Contrib. Amer. Ent. Inst., vol. 20, 1983

The nominate subspecies (with hind tarsus yellowish-white or brownish-

yellow) occurs in Vietnam and Japan. The subspecies

(Turner) (with blackish hind tarsus) occurs in North China.

Mandibles evenly curved (Figs. 22, 31, 65); 2nd tergite smooth; apex of Ist

tergite (partly) smooth (Figs. 53, 70); length of ovipositor sheath 1.24 -

1.58 (exceptionally 1.79) times fore wing; occipital carina arched

medio-dorsally (Figs. 21, 52) or reduced (Fig. 90); length of hind tibia

1.6 -1.9 times hind femur; clypeus more or less convex or margin pro-

truding; (Brulleia Szépligeti) Sc eerie esi eee 2 ee Se ka ee

2. Labial palp with 3 segments (Figs. 20, 36); maxillary palp with 4 or 5

segments (Figs. 20, 25, 54); marginal cell of hind wing narrower

apically, its apical width 2.3 -3.0 (exceptionally 4.0) times the minimum

width of cell below vein RI (Figs. 17, 50); notauli comparatively wide

aoe Visine? sippecteriony (Fics. 23,°95)". 8 ee ob ee 3

Labial palp with 2 segments (Figs. 62, 77); maxillary palp with 2 or 3 seg-

ments (Figs. 77, 89), exceptionally 4th segment faintly indicated basally

(Fig. 62); marginal cell of hind wing rather wide apically, its apical

width 3-4 times the minimum width of cell below vein RI (Figs. 61, 87);

oe comparatively narrow subposteriorly (Fig. 69) or reduced (Fig.

a ee Ce ee oe eee ee 6

3. Maxillary palp with 5 segments (Fig. 20) and its length 0.5 -0.7 times

height of head (Fig. 15); prepectal carina complete (Fig. 15) ......

a a a a Se Be ews Se 2. nipponensis,;, nN. sp.

Maxillary palp with 4 segments (Figs. 25, 36) and its length ca. 0.3 times

height of head (Fig. 24); prepectal carina reduced anteriorly (Fig. 24)

and medio-ventrally obsolescent or absent (Fig. 35) . 2.0.2... - 4

4. Antenna with no whitish band; pronope short, scarcely wider than long (Fig.

33); dorsal half of antenna yellowish; Ist tergite robust, its length ca.

L4 times its apical width (Pig. 29). 2" ok. 8, townes?, nl sp:

Antenna with whitish band; pronope strongly transverse, much wider than

long (Figs. 42, 55); basal half of antenna largely blackish; Ist tergite

slender, its length 2.1-4.6 times its apical width (Figs. 46, 53) ... 5

5. Body completely brownish-yellow; basal half of vein 2A of fore wing not

sclerotized (Fig. 37); bases of fore and middle tibiae yellowish; labrum

subtmineate apically (Pig. 40). 2 Sere Pee 4. “brannea;, nn. sp.

Body almost completely black; basal half of vein 2A of fore wing

sclerotized (Fig. 50); bases of fore and middle tibiae whitish; labrum

geenly. emartingte (Fig, 56). 2 cee ee eh 5, 8igfa, fh. Sp.

6. Maxillary palp with 3 segments, 4th segment more or less faintly indicated

(Fig. 62); prepectal carina complete and strong (Fig. 59); Ist tergite

slender (Figs. 67, 70), its length 2.1-4.6 times its apical width; dorsal

carinae of Ist tergite distinct basally (Fig. 70). 6. tricolor, on. sp.

Maxillary palp with 2 segments (Figs. 77, 89); prepectal carina reduced

dorsally (Fig. 71) and weak (Fig. 84); Ist tergite usually rather robust

(Figs. 83, 98), its length about 1.8 (exceptionally about 3) times its

apical width; dorsal carinae of Ist tergite reduced basally (Figs. 83)

Of GUSCHt Vee ee oa ee ee Re ea 7

van Achterberg: Brulleiini (Braconidae) 287

7. Body (except head) brownish-yellow, without metallic sheen; base of hind

tibia and whole hind tarsus yellowish or brownish; clypeus flat medially

or nearly so (Fig. 73); 15th and 16th antennal segments dark brown or

DIGG oy sgl i es ea ne eee 7. melanocephala Szépligeti

Body largely black with a bluish metallic sheen; base of hind tibia and

hind tarsus (except telotarsus) white; clypeus medially distinctly de-

pressed (Fig. 91); Ist and 16th antennal segments white ........ :

yee aes ee .8. latiannulata (Cameron), n. comb.

PARABRULLEIA, n. genus

Type-species: Doryctes shibuensis Matsumura, 1912.

Etymology: The word "para" (Greek for "near") is added to the name

Brulleia, because of the new genus is closely related to the genus Brulleia.

Gender: feminine.

Mandibles angularly bent medially (Fig. 6); maxillary and labial palpi

with 4 and 3 segments, respectively (Fig. 8); clypeus flat (Fig. 6); occipital

carina evenly curved, not arched (Fig. 9); vertex without distinct medial groove;

frons medially distinctly concave (Fig. 6); length of hind tibia 2.3 -2.4 times

hind femur; 2nd tergite sculptured basally (Fig. 12).

Distribution: East Palearctic and North East Oriental regions; one

species.

1. Parabrulleia shibuensis (Matsumura), n. comb. (Figs. 2-14, 100)

Doryctes shibuensis Matsumura, 1912: 151], pl. 52-1.

Brulleia shibuensis: Shenefelt, 1970: 190; Watanabe, 1972: 7, Figs, 5, 6.

Brulleia chinensis Turner, 1918: 171; Shenefelt, 1970: 190.

Brulleia euphemia Turner, 1918: 387; Shenefelt, 1970: 190. Syn. nov.

Redescribed from the holotype of Brulleiaeuphemia Turner, female.

Length: Body 22 mm. Fore wing 17.1 mm.

Head: Antennal segments 33, but apical segments missing, length of 3rd

segment |.] times 4th segment, length of 3rd and 4th segments 4.2 and 3.9

times their width, respectively; annellus scarcely differentiated (Fig. 10);

length of maxillary palp 0.3 times height of head; length of eye in dorsal view

1.2 times temple; temple punctulate (Fig. 9); POL : Mocellus : OOL = 4: 5

: 10; frons rugose, but punctulate near eyes (Fig. 9); face coarsely reticulate-

rugose; clypeus shallowly punctate (Fig. 6); malar space coriaceous, its length

0.7 times basal width of mandible.

Mesosoma: Length of mesosoma 1.7 times its height; pronope very large,

wide, spindle-shaped (Fig. 13); side of pronotum medially and posteriorly

coarsely crenulate, ventrally punctulate, rest smooth (Fig. 2); prepectal carina

rather weak and irregular, medio-ventrally absent; precoxal sulcus complete,

punctate-rugose (Fig. 2); notauli deep, complete and widely crenulate (Fig.

13); scutellum rather flat and punctulate; propodeum coarsely reticulate, but

antero-laterally coriaceous (Fig. 13).

288 Contrib. Amer. Ent. Inst., vol. 20, 1983

Wings: Fore wing! 1 :°3-SR.: SRI =.9 2:12. :.43; d-CUl 2. 2+CUl a3): 32;

2-SR : 3-SR : r-m = 9: 12: 12. Hind wing: cu-a sinuate and strongly in-

clivous (Fig. 3).

Legs: Length of femur, tibia and basitarsus of hind leg 5.1, 17.8, and 10.7

times their width, respectively; length of hind tibia 2.3 times hind femur (Fig.

14).

Metasoma: Length of Ist tergite 2.0 times its apical width, its surface

coarsely reticulate medially, laterally and basally largely crenulate (Fig. 12),

dorsal carinae present in front of spiracles; length of ovipositor sheath 1.91

times fore wing.

Color: Yellowish-brown; antenna (except two basal segments, annellus,

and 10th -l6th segments) and ovipositor sheath, black; fore wing anteriorly (in-

cluding pterostigma) and apices of 3rd-8th tergites, 5th sternite and hypo-

pygium partly, and middle of mesoscutal lobes, more or less dark brown; rest

of wing veins brown; 10th -l6th antennal segments and middle and hind tarsi,

yellowish-white; wing membrane brown.

Holotype of euphemia in British Museum (Natural History), London: "Type,

H. T.", "B. M. Type, Hym., 3.c.898", "Brulleia euphemia Turn., Type",

"Tonkin, V. 1917, R. V. de Salvaza", "Indo-China, R. V. de Salvaza, 1919-25".

Additionally examined: 1) Holotype of Doryctes shibuensis Matsumura from

Japan (Entomological Institute, Sapporo; F); agrees with holotype of euphemia.

but the hind tarsus is yellowish-brown. 2) Holotype of Brulleia chinensis

from North China [British Museum (Natural History); M], hind tarsus and

apical half of hind tibia, blackish; Ist tergite densely vermiculate and coarsely

rugose; basal 0.7 of 2nd tergite largely and distinctly rugose; vein r-m of fore

wing scarcely bent (Fig. 100); malar space rugose. 3) 4 F from Japan: Tokyo

(Townes Collection, Ann Arbor; Entomological Institute, Sapporo), Odawara

(Kanagawa Pref., Honshu), and Mt. Hiko (Fukuoka Pref., Kyushu). Both lat-

ter specimens in Kyushu University Collection, Fukuoka.

Variation: Antennal segments 41 (1 F); length penultimate segment ca.

1.4 times its width; length of fore wing 17.1-19.9 mm, of body 22-24.8 mm;

length of ovipositor sheath 1.6 ~-1.91 times fore wing; marginal cell of hind wing

usually wider apically than figured; length of hind tibia 2.3 -2.4 times hind

femur.

Genus BRULLEIA Szépligeti

Type-species: Brulleia melanocephala Szépligeti, 1904.

| Diagnosis: Mandibles evenly curved (Figs, 22, 31, 65); maxillary and labial

palpi 2-5 and 2-3 segments, respectively (Figs. 20, 36, 62, 77); face densely

reticulate-rugose; clypeus more or less convex or medially depressed (Figs. 22,

40, 56, 65); occipital carina arched medio-dorsally (Figs. 21, 30, 52) or reduced

(Fig. 90); vertex usually with longitudinal groove (Figs. 21, 44); frons weakly

concave medially (Figs. 64, 75, 90) or nearly flat (Figs. 44, 52); length of

hind tibia 1.6 -1.9 times hind femur; 2nd tergite smooth.

Distribution: East Palearctic, East Oriental and New Guinea; seven

species.

M = male. F = Female.

van Achterberg: Brulleiini (Braconidae) 289

2. Brulleia nipponensis, n. sp. (Figs. 15 -23, 92 -96)

Brulleia euphemia: Watanabe, 1972: 7 (nec Turner, 1919).

Length: Body 16.1 mm. Fore wing 12.4 mm.

Head: Antennal segments 41, length of 3rd segment 1.3 times 4th segment,

length of 3rd and 4th segments 5.0 and 4.0 times their width, respectively,

the penultimate segment quadrate (Fig. 18); maxillary and labial palpi with

5 and 3 segments, respectively (Fig. 20); length of maxillary palp 0.6 times

height of head; length of eye in dorsal view 1.1 times temple; temple punctate

(Fig. 21); POL: @& ocellus : OOL = 6: 5 : 13; frons largely curved striate (Fig.

21); clypeus coarsely punctate (Fig. 22); malar space rugose, its length 0.9

times basal width of mandible.

Mesosoma: Length of mesosoma 1.6 times its height; pronope medium -

sized and slit-shaped (Fig. 23); side of pronotum rugose except for the punc-

tulate dorsal part and the crenulate medial groove (Fig. 15); prepectal carina

complete; precoxal sulcus complete, crenulate-rugose and area below it densely

punctate (Fig. 15); notauli deep, widely crenulate posteriorly (Fig. 23); scutellum

rather convex and densely punctate; propodeum reticulate-rugose, but antero-

laterally punctate, rugae more widely spaced than in shibuensis.

Wings: Fore wit: #1, 3-8R: + SR) «75 Te 435 CUP 2-CUl ee 1s 123

2-SR : 3-SR : r-m = Il: ll: 14. Hind wing: cu-a inclivous and posteriorly

curved towards wing base (Fig. 17).

Legs: Length of femur, tibia and basitarsus of hind leg 5.8, ll.2 and 9.4

times their width, respectively; length of hind tibia 1.6 times hind femur (Fig.

95).

Metasoma: Length of Ist tergite 1.8 times its apical width, its surface

rather sparsely rugose and posteriorly smooth (Fig. 96); dorsal carinae present

in front of spiracles; length of ovipositor sheath 1.4] times fore wing.

Color: Brownish-yellow; mandibles, pterostigma (but basally and apically

narrowly yellowish), wing veins, antenna (but Ist and 2nd segments brownish-

yellow and 10th -1l4th segments yellowish-white), and ovipositor sheath, dark

brown or blackish; apex of hind tibia slightly infuscated; wing membrane yellow-

ish.

Holotype: Female, Fukuoka: Mt. Kanayama, Fukuoka Pref. (Kyushu,

Japan), ll. viii. 1979, R. Noda leg. Type no. 2229. (Entomological Laboratory,

- Kyushu University). Paratype: 1 F, Hirakura, Mie Honshu, 17. viii. 1958,

M. Matsuura (Rijksmuseum van Natuurlijke Historie, Leiden). 1 F, Sata,

Kagoshima, 24. iv. 1964, Y. Miyake. 1 F, Zyozankei, Hokkaido, 17. ix. 1954,

T. Kumata (Entomological Institute, Sapporo).

Variation: Length of fore wing 10.1 -13.5 mm, of body 12.2 -18.2 mm;

length of maxillar palp 0.5 -0.7 times height of head; length of ovipositor

sheath 1.41 -1.56 times fore wing.

3. Brulleia townesi, n. sp. (Figs. 24 -34)

Length: 11.7 mm, fore wing 9.7 mm.

Head: Antennal segments 41, length of 3rd segment 1.3 times 4th segment,

length of 3rd and 4th segments 3.5 and 2.8 times their width, respectively,

290 Contrib. Amer. Ent. Inst., vol. 20, 1983

length of penultimate segment 0.9 times its width (Fig. 27); maxillary and

labial palpi with 4 and 3 segments, respectively (Fig. 25); length of maxil-

lary palp 0.3 times height of head; length of eye in dorsal view 1.5 times

temple; temple punctate (Fig. 24); POL: ® ocellus: OOL = 9: 7 : 23; frons

medially largely smooth, laterally striate (Fig. 30); clypeus punctate (Fig. 31);

malar space rugose, its length 0.8 times basal width of mandible.

Mesosoma: Length of mesosoma I.5 times its height; pronope large and

transverse (Fig. 33); side of pronotum ventrally largely smooth, medially

crenulate, rest partly rugose-crenulate (Fig. 24); prepectal carina laterally

reduced, weak and medio-ventrally absent; precoxal sulcus complete, reticulate -

rugose (Fig. 24); notauli rather deep and narrowly crenulate, but posteriorly

wider (Fig. 33); scutellum convex, densely and finely punctate (Fig. 33); pro-

podeum medially transversely rugose, laterally finer rugose.

Wings: Fore wing: r: 3-SR:: SRI = 7: ll : 40; cu-a subinterstitial;

2-SR : 3-SR.: r-m = 10: 11 : 10. Hind wing: cu-a inclivous and only slightly

bent posteriorly (Fig. 26).

Legs: Length of femur, tibia and basitarsus of hind leg 4.6, 12.3, and 10.4

times its width, respectively; length ofhind tibia 1.7 times hind femur (Fig.

32). |

Metasoma: Length of Ist tergite 1.4 times its apical width, its surface

medially smooth, laterally rugose (Fig. 29); dorsal carinae weak in front of

spiracles; length of ovipositor sheath 1.58 times fore wing.

Color: Brownish-yellow; basal 17 segments of antenna whitish-yellow, rest

dark brown; pterostigma, wing veins and ovipositor sheath, dark brown; side

of mesosoma, stemmaticum and apex of Ist tergite with faint infuscated

patches; wing membrane light brown; legs (including hind tarsus) light yellowish.

Holotype: Female, Philippines: Mindoro, Alcate, Victoria, iv. ll. 1954, H.,

M. & D. Townes (Townes).

4. Brulleia brunnea, n. sp. (Figs. 35-46)

Length: Body 12.2 -13.1 mm. Fore wing Il.1 mm.

Head: Antennal segments 43 -44, length of 3rd segment 1.2 times 4th

segment, length of 3rd and 4th segments 4.2 and 3.6 times their width, res-

pectively; the penultimate segment quadrate (Fig. 38); maxillary and labial

palpi with 4 and 3 segments, respectively (Fig. 36); length of maxillary palp

0.3 times height of head; length of eye in dorsal view 1.3 times temple;

temple finely punctate (Fig. 35); POL : & ocellus : 0O0L = 9: 7: 18; frons

rugose (Fig. 44); clypeus rugose (Fig. 40); labrum truncate ventrally (Fig. 40);

malar space rugose, its length 0.9 times basal width of mandible.

Mesosoma: Length of mesosoma 1.5 times its height; pronope very wide,

slit-shaped and deep (Fig. 42); side of pronotum largely smooth, but medio-

anteriorly crenulate and rugose-punctate ventrally (Fig. 35); prepectal carina

only laterally largely present (Fig. 35); precoxal sulcus complete, strongly

rugose; notauli complete, posteriorly widely rugose-crenulate (Fig. 42);

van Achterberg: Brulleiini (Braconidae) 291

scutellum rather flat, densely and finely punctate; propodeum coarsely

reticulate-rugose, except anterolaterally.

Wings’, Pore witg:) 12 3-5R Skil ='7 1) 7 4 BCUl ? 220Ul = 3°. 3:

Z2-SR : 3-SR : r-m = 19 : 30: 20; 2A largely unsclerotized. Hind wing:

cu-a inclivous, somewhat sinuate (Fig. 37).

Legs: Length of femur, tibia and basitarsus of hind leg 6.4, 15.2 and 10.8

times its width, respectively; length of hind tibia 1.8 times hind femur (Fig.

45).

Metasoma: Length of Ist tergite 2.1-2.4 times its apical width, its sur-

face rather coarsely reticulate-rugose, but basally and apically narrowly smooth

(Fig. 46), dorsal carinae obsolete in front of spiracles; length of ovipositor

sheath 1.46 -1.54 times fore wing.

Color: Brownish-yellow; hind tarsus (except base), and llth (except base)

19th (except apex) antennal segments, white; rest of antenna (except scapus,

pedicellus and radix), vein C+SC+R (except base), pterostigma, and ovipositor

sheath, black or blackish-brown; rest of wing veins dark brown; wing membrane

light brown; humeral plate whitish; apex of hind tibia slightly infuscated.

Holotype and paratype: Female, Java: G. Tangkoeban Prahoe, 4000 - 5000

ft., Preanger, Ill. 1938. Paratype: 1 F, topotypic, v. 1937, F. C. Drescher

(Rijksmuseum, Leiden).

5. Brulleia nigra, n. sp. (Figs. 47 -58)

Length: Body 14.6 mm. Fore wing Il.7 mm.

Head: Antennal segments 44, length of 3rd segment 1.2 times 4th segment,

length of 3rd and 4th segments 4.1 and 3.4 times their width, respectively,

length of penultimate segment 0.9 times its width (Fig. 49); maxillary and

labial palpi with 4 and 3 segments, respectively (Fig. 54); length of maxillary

palp 0.3 times height of head; length of eye in dorsal view 1.4 times temple;

temple finely rugose, laterally striate (Fig. 52); clypeus rugose (Fig. 56);

labrum emarginate ventrally (Fig. 56); malar space rugose, its length 0.7

times basal width of mandible.

Mesosoma: Length of mesosoma 1.6 times its height; pronope very wide,

deep and, elliptical (Fig. 55); side of pronotum medio-dorsally crenulate,

ventrally striate, posteriorly punctate, and rest smooth (Fig. 47); prepectal

carina laterally weak and ventrally absent (Fig. 47); precoxal sulcus complete

_ and rugose (Fig. 47); notauli complete and posteriorly widely crenulate (Fig.

55); scutellum rather convex and sparsely punctulate (Fig. 55); propodeum

largely reticulate-rugose and antero-laterally smooth.

Wings:., Pore wing: (f° 3.3-SRia--BRI. 8. 18+ 4 26.2. 38) d-CU i 2-OU) = 4

10; 2-SR : 3-SR : r-m = 19 : 26: 14; 2A distinctly sclerotized basally. Hind

wing: cu-a inclivous, somewhat bent posteriorly (Fig. 50).

Legs: Length of femur, tibia and basitarsus of hind leg 6.7, 13.7, and

11.0 times their width, respectively; length of hind tibia 1.8 times hind femur

(Fig. 57).

Metasoma: Length of Ist tergite 2.1 times its apical width, its surface

rugose, but basally and apically smooth (Fig. 53), dorsal carinae absent; length

of ovipositor sheath 1.44 times fore wing.

292 Contrib. Amer. Ent. Inst., vol. 20, 1983

Color: Black; Radix, scapus, pedicellus, and annellus of antenna, fore leg

(except basal 0.7 of fore tibia), middle coxa, metasoma ventro-basally, apex

of 3rd tergite, 8th tergite, wing veins largely, pterostigma, and humeral plate,

(dark) brown; hind tarsus (except base), Ilth-1l6th antennal segments, white;

palpi, yellowish; basal 0.7 of fore tibia, basal 0.3 of middle tibia, 4th and 5th

middle tarsal segments, whitish; rest of middle leg, hind coxa and trochanters;

blackish-brown. |

Holotype: Female, Java: Tjibodas, 1400 m, 27-29. vii. 1930, Lieftinck

(Rijksmuseum van Natuurlijke Historie, Leiden).

6. Brulleia tricolor, n. sp. (Figs. 59 ~70)

Length: Body 11.8 mm. Fore wing 10.5 mm.

Head: Antennal segments 40, length of 3rd segment 1.2 times 4th segment,

length of 3rd and 4th segments 4.7 and 4.0 times their width, respectively,

length of penultimate segment 1.2 times its width (Fig. 60); annellus differen-

tiated (Fig. 59); maxillary and labial palpi with 3, (but 4th faintly indicated)

and 2 segments, respectively (Fig. 62); length of maxillary palp 0.2 times

height of head; length of eye in dorsal view 1.3 times temple; temple punctu-

late (Fig. 59); POL: &@ ocellus : OOL - 7: 4: 12; frons largely smooth medial-

ly, laterally striate (Fig. 64); clypeus rugose (Fig. 65); malar space rugose,

its length 0.7 times basal width of mandible.

Mesosoma: Length of mesosoma 1.5 times its height; pronope wide, deep

and spindle-shaped (Fig. 69) side of pronotum medially crenulate, rest largely

smooth (Fig. 59); prepectal carina complete and lamelliform (Fig. 59). Pre-

coxal sulcus complete and rugose; notauli complete, narrowly crenulate post-

eriorly (Fig. 69); suctellum rather convex and punctulate. Propodeum reticu-

late, but anteriorly largely smooth and with short medial carina (Fig. 69).

WES) Ate Winks: Tossaak come 2.3. 3 IOs Cul e 26CUL ue. 2 2.17

2-SR : 3-SR : r-m = Il: 12: 10. Hind wing: cu-a inclivous and distinctly

sinuate (Fig. 65).

Legs: Length of femur, tibia and basitarsus of hind leg of F-paratype

(missing in holotype) 5.9, 15.7, and 11.6 times its width, respectively; length

of hind tibia 1.8 times hind femur (Fig. 66).

Metasoma: Length of Ist tergite 2.3 times its apical width, its surface

behind spiracels finely reticulate-rugose, but medially, posteriorly and in front

of spiracles smooth (Fig. 70); dorsal carinae present in basal 0.6 of Ist ter-

gite; length of ovipositor sheath 1.54 times fore wing.

Color: Black; coxae, trochanters, femora, metanotum, propodeum, Ist and

2nd tergites, reddish-brown; base of femora, middle basitarsus [and hind tarsus

(except for brownish telotarsus) of paratypes], and 10th -17th antennal segments

white; fore tarsus, rest of middle tarsus and wings, infuscated; scapus, pedicel-

lus, apical 0.7 of fore and middle tibiae and pterostigma, dark brown.

Holotype: Female, Philippines: Mt. Canlaon, 3600 ft., Negros Or., May

8, 1953, H., M. & D. Townes (Townes). Paratypes: | F and 3 M, #1F,

topotypic (Rijksmuseum van Natuurlijke Historie, Leiden). 2 M, topotypic,

April 29, 1953 and May 8, 1953, (Townes) Collection. 1 M, Illong, Mt. Halcon,

4500 ft., Mdro. Or., v. ll. '54, Phil., M. & D Townes (Townes).

van Achterberg: Brulleiini (Braconidae) ; 293

Variation: Length of fore wing 10.5 -10.6 mm (F) or 6.8-9.5 mm (M),

of body 11.8-12.8 mm (F) or 9.0-13.5 mm (M); antennal segments of M 35-

38; length of Ist tergite 2.1-2.3 times its apical width (F) or 3.7 -4.6 times

(M, Fig. 67); M-paratypes have only llth ~13th antennal segments white, hind

basitarsus brown or white basally, 4th maxillary palp segment in 1 M almost

completely differentiated, and mesosoma (except for Ist tergite) brown.

7. Brulleia melanocephala Szépligeti (Figs. 71-83)

Brulleia melanocephala Szépligeti, 1904: 150, Fig. 52. Shenefelt, 1970: 190.

Cenocoelius annulicornis Cameron, 191]: 245-247; Shenefelt, 1970: 180;

Van Achterberg, 1980: 210. n. syn.

Length: Body 14.8 mm. Fore wing 11.5 mm.

Head: Antennal segments 41, length of 3rd segment 1.1 times 4th segment,

length of 3rd and 4th segments 4.8 and 4.4 times their width, respectively,

length of penultimate segment 1.2 times its width (Fig. 72); annellus distinct

(Fig. 76); maxillary and labial palpi both with 2 segments (Fig. 77); maxillary

palp not or slightly protruding below head (Fig. 71); eye in dorsal view 1.6 times

temple; temple smooth (Fig. 71); POL : & ocellus : OOL = ll : 9 : 24; frons

with some stria, largely smooth (Fig. 75); clypeus coarsely reticulate (Fig. 73);

malar space rugose, its length 0.9 times basal width of mandible.

Mesosoma: Length of mesosoma 1.5 times its height; pronope spindle-shaped,

deep and wide; side of pronotum smooth, but medio-anteriorly and posteriorly

crenulate (Fig. 71); prepectal carina absent laterally (Fig. 71), ventrally pres-

ent; only posterior half of precoxal sulcus rugose, rest largely smooth and shal-

low (Fig. 71); notauli complete, largely smooth, and narrow posteriorly (Fig.

82); scutellum convex and punctulate; propodeum coarsely reticulate-rugose,

with a short medial carina anteriorly.

Wings: Fore wing? (7 :°3-SR BRP 6 Oy Bes oye1eCUl + 20 hs

22; 2-SR : 3-SR : r-m = 13 : 18 : 16. Hind wing: cu-a inclivous and rather

curved (Fig. 79).

Legs: Length of femur, tibia and basitarsus of hind leg 4.9, 13.6, and

13.2 times their width, respectively; length of hind tibia 1.8 times hind femur

(Fig. 78).

Metasoma: Length of Ist tergite 1.8 times its apical width, its surface

largely smooth, except for some rugae behind spiracles (Fig. 83); length of

- ovipositor sheath 1.39 times fore wing.

Holotype in Budapest Museum: New Guinea: Biro, iv. 4. 1901, Mt.

Hansemann, Astrolabe B., 1 F (Holotype), "Brulleia melanocephala Szépl.,

1904, det. Papp., 1967", "Hym. Typ. No. 782, Mus. Budapest". Holotype of

Cenocoelius annulicornis Cameron (Instituut voor Taxonomische Zoologie,

Amsterdam): New Guinea: Lorentz, 1909-10, Bivak Eiland, ix. 09,

"Cenocoelius annulicornis Cam., Type" (in Cameron's handwriting). Further

1 F examined (Townes Collection, Ann Arbor): Baiyer R., (Papua) N. Guinea,

II. 6-25. 1979, 1100 m, J. Sedlacek.

Variation: Length of fore wing 11.3 -12.4 mm, of body 14.0 -15.5 mm;

antennal segments 41-43; length of ovipositor sheath 1.24 -1.51 times fore wing;

the holotype of annulicornis has posterior half of Ist tergite extensively

294 Contrib. Amer. Ent. Inst., vol. 20, 1983

sculptured; wing membrane infuscated, and precoxal sulcus completely sculptured;

the F from Papua New Guinea has scapus and pedicellus dark brown or blackish

and only Ilth-13th antennal segments whitish.

8. Brulleia latiannulata (Cameron), n. comb. (Fig. 84-91, 97-99, 101)

Cenocoelius latiannulatus Cameron, 191]: 236; Shenefelt, 1970: 181; Van

Achterberg, 1980: 211.

Length: Body 17.5 mm. Fore wing 13.0 mm.

Head: Remaining antennal segments 21, apical segments missing, length

of 3rd segment 1.2 times 4th segment, length of 3rd and 4th segments 5.2 and

4.2 times their width, respectively; annellus distinct (Fig. 85); maxillary and

labial palpi both with 2 segments (Fig. 89); maxillary palp slightly protruding

below head (Fig. 84); eye in dorsal view 1.1] times temple; temple largely

smooth (Fig. 84); occipital carina reduced medio-dorsally (Fig. 90); POL : &

ocellus : OOL = 12 : 6: 6; frons with some striae (Fig. 90); clypeus reticulate-

rugose (Fig. 91); malar space rugose, its length 0.8 times basal width of man-

dible.

Mesosoma: Length of mesosoma 1.5 times its height; pronope, spindle-

shaped, deep and large; side of pronotum medially smooth, rest crenulate or

rugulose (Fig. 84); prepectal carina nearly complete, weak behind fore coxae;

precoxal sulcus complete, coarsely rugose; notauli complete, posteriorly narrow-

ly crenulate (Fig. 101); scutellum convex and punctulate; propodeum coarsely

reticulate, but anteriorly smooth.

Wings: Fore wing? rss 3+SR.:: SRE =: 8 2.12 4-40; 1-CUl. 2: 2-CUL =:5.:. 32;

2-SR : 3-SR : r-m = 10: 12 : 12. Hind wing: cu-a inclivous and sinuate (Fig.

87).

Legs: Length of femur, tibia and basitarsus of hind leg 5.1, 14.3 and 13.2

times their width, respectively; length of hind tibia 1.9 times hind femur (Fig.

97 )«

Metasoma: Length of Ist tergite 1.8 times its apical width, its surface

behind spiracles coarsely reticulate-rugose (Fig. 98), dorsal carinae absent;

length of ovipositor sheath 1.79 times fore wing.

Color: Black with a bluish metallic sheen; 10th -l6th antennal segments,

basal third of hind tibia, hind tarsus, trochanters (largely), middle tibia (except

apex), fore tibia, more or less whitish; scapus, pedicellus, annellus, palpi, man-

dibles, pterostigma, wing veins, tegulae, more or less dark brown; fore tarsus,

fore femur dorsally, and middle tarsus (except basitarsus), brownish; epipleuron

of Ist tergite, apical part of 2nd and basal margin of 3rd epipleura, whitish-

yellow; wing membrane rather infuscated, more strongly near veins r and 3-

SR, and pterostigma of fore wing (Fig. 86).

Holotype in Instituut voor Taxonomische Zoologie, Amsterdam: New

Guinea: Lorentz, 1909-10, Bivak Ejiland, I. 10,"Cenocoelius latiannulatus Cam.,

Type" (in Cameron's handwriting). Additionally examined 4 F (3 in Townes

Collection, Ann Arbor; | in Rijksmuseum van Natuurlijke Historie, Leiden) from

Papua New Guinea. Collected in Saruwaged Mountains (500 m, I. 22-Il. 16.

1979), Laea Zdnag Road (200 m, I. 14-19. 1979), Busu River, 60 km East of

Lae, 20 m, I. 13 -IIl. 10. 1979), and Gerit River (650 m, Jimmi Valley, II. 7 -

26. 1979).

van Achterberg: Brulleiini (Braconidae) 295

Variation: Length of fore wing 7.3 -13.0 mm, of body 9.0 -17.5 mm;

antennal segments 39-41; length of Ist tergite 1.8 -3.1 times its apical width,

surface sometimes largely smooth; length of ovipositor sheath 1.56 -1.79 times

fore wing; scapus blackish or dark brown; occipital carina obsolete and arched

dorsally or medially absent.

ACKNOWLEDGMENTS

I wish to express my gratitude to the following persons for providing type-

specimens, unidentified specimens, or valuable comment: Mr. T. Huddleston

(London); mr. K. Maeto (Fukuoka); Dr. P. Oosterbroek (Amsterdam); Dr. J.

Papp (Budapest); Drs. M. Suwa and C. Watanabe (Sapporo); and Drs. H. K. and

M. Townes (Ann Arbor).

REFERENCES

Achterberg, C. van. 1979. A revision of the subfamily Zelinae auct. (Hym.,

Braconidae). Tijdschr. Ent. 122: 241-479, Figs. 1-900.

Achterberg, C. van. 1980. The Cameron types of Braconidae in the Nether-

lands (Hym., Ichneumonoidea). Bull. Zool. Mus. Univ. Amst. 7: 209-214.

Cameron, P. 1911. Hymenoptera (except Anthophila and Formicidae). Nova

Guinea 9: 185-248.

Matsumura, S. 1912. Thousand insects of Japan, Suppl. 4: 1-247. Tokyo.

Shenefelt, R. D. 1970. Braconidae, pt. 2. Hymenopterorum Catalogues (nov.

ed.) 5: 177-306.

Szépligeti, G. V. 1904. Fam. Braconidae. Genera Insectorum 22: 1-253,

Figs. 1-32.

Turner, R. E. 1918. Notes on the Braconidae in the British Museum. IV.

On new Helconinae, mostly Australian. Ann. Mag. Nat. Hist. (9)2: 163-

173;

Turner, R. E. 1919. On Indo-Chinese Hymenoptera collected by R. Vitalis de

Salvaza, ili. Ann. Mag. Nat. Hist. (9)4: 384-394.

Watanabe, C. 1972. A revision of the Helconini of Japan and a review of the

Helconine genera of the world (Hym., Braconidae). Insecta Matsumurana

35:. 1418, Figs. 1-8.

Contrib. Amer. Ent. Inst., vol. 20, 1983

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ON EVOLUTION OF ENDOPARASITISM: THE BIOLOGY OF SOME

GENERA OF ROGADINAE (BRACONIDAE)

Mark R. Shaw

Royal Scottish Museum, Edinburgh, U.K.

INTRODUCTION

Both ectoparasitism and endoparasitism are found in several families of

parasitic Hymenoptera (parasitoids) and, since the former is generally accepted

as being the more primitive, it is evident that there have been many independ-

ent evolutions of endoparasitism from different ectoparasitic ancestral stocks.

One group in which this has occurred is the braconid subfamily Rogadinae

(sensu Achterberg 1976), in which elements of the extant tribes Exothecini

(ectoparasitic) and Rogadini (endoparasitic) are clearly closely related. I have

investigated the biology of British species of some of the genera in these two

tribes, considering particularly host range, the biology of the hosts, the host

stages attacked, the use and effects of parasitoid venom, and egg placement.

From this it has been possible to trace developments in the ectoparasitic habit,

as efficient exploitations of given host life-histories, to the point at which

it becomes so specialized and refined that it differs from the most primitive

condition of endoparasitism I have seen in the Rogadini only in the relatively

minor detail of egg placement on different sides of the host's integument.

The meaning of host range. Most parasitoids are not absolutely specific

to single host species but instead tend to attack a group of hosts which usually

bear some relation, either ecological or phylogenetic, to one another. The

"host range" (or ‘usual hosts") of a given parasitoid is a key feature of its

biology, although in practice it is a nebulous term difficult to define with any

precision. This is because some host species may be attacked less avidly, or

with less success, than others, and furthermore it is known that such factors

can vary from one population of a parasitoid species to another: indeed, some

plasticity in behavior and variability in success have presumably been pre-

requisites in the evolution of host specializations at all. However, it is useful

to define host range in general terms as including only the species of potential

hosts that the parasitoid is usually able to attack successfully, following a

pattern of searching behavior enabling it to encounter them regularly. Poten-

tial hosts that are attacked only by some freak of circumstance, or which

only occasionally permit the parasitoid's full development, would thus be ex-

cluded from a useful concept of host range, as would species of otherwise suit=

able hosts which the parasitoid rarely encounters. In practice host ranges can

usually be understood as a balance - more or less severe, and variously biased -

between host ecology and host phylogeny. Not all the elements of a host

range will necessarily be present at a particular site, however, so at a local

level parasitoid populations often attack only a more limited set of hosts.

Also, a particular host species may not be available during the whole span of

a parasitoid's period of activity, especially if, as is often the case in temperate

areas, the parasitoid is plurivoltine and some or all of its hosts are univoltine.

A regular parasitoid of a given host must first search in the environment

supporting the host, second be able to detect and accept the host, and third

be physiologically capable of full and unimpaired development upon it. There

are thus two broad aspects of host range involving potential behavioral

307

308 Contrio. Amer. Ent. inst., vol. 20, 1983

flexibility: (a) host habitat selection, or where to search, and (b) host selec-

tion, or what to attack. Host habitat selection is one of the simplest expres-

sions of specialization, and very many parasitoids of phytophagous hosts search

exclusively either on arborescent vegetation or in the field layer. The im-

portance of the second kind of behavioral limitation to host range, whereby

certain potential host species are usually rejected although proving to be suit-

able on the rare occasions they are attacked, is often seen in ectoparasites,

which are in general under less severe physiological pressures than typical

endoparasites and therefore tend to have broader potential host compatibilities.

MATERIALS AND METHODS

As wide a range as possible of larvae Lepidoptera in general and leaf-

mining insects of the Orders Lepidoptera, Hymenoptera, Coleoptera and, to

a lesser extent, Diptera have been collected wild from a diverse range of the

habitats available in Britain and reared in captivity, firstly to obtain host

records of parasitoids in general, and secondly to obtain culture material for

experimental rearings. Except that some experiments involving cultures of

leaf-miners and their parasitoids were conducted in muslin sleeves on trees,

all material was reared in an unheated detached wooden shed, which was

situated in deep shade, painted white, and permanently ventilated by replacing

the window and door with wire mesh. Under these conditions air tempera-

tures were always within 1°C of outdoor shade, and natural conditions of

humidity and light were maintained at least approximately: in all cases that

could be checked, emergence dates of captive host and parasitoid species cor-

responded to those in the field. Wild collected larvae and experimental cul-

tures were kept as single species in carefully counted groups (of often indi-

vidually) in closed tubes or plastic boxes. All larval hosts were fed on

natural foodplants: leaf-mining species unable to change leaves were mostly

collected as mature (or already killed or vacated) entities and the mines were

cut out and tubed, or otherwise processed, individually.

In addition to my own rearings of parasitoids, | have received much rear-

ed material from lepidopterists and other entomologists who, by screening so

large a number of potential hosts, have guided me to hosts producing

Rogadinae on a number of occasions. Among several entomologists, B. T.

Parsons, R. A. Softly and the late W. A. Watson have been particularly gener-

ous with their time spent searching for and collecting wild host species sus-

pected of harboring Rogadinae. All of this has helped me to at least orien-

tate towards the ideal of sampling the natural populations of parasitoids to

find out what their hosts are, rather than merely sampling host populations

to see what their parasitoids are.

At least some of the experimental parasitizations were carried out under

full observation for all species of Rogadinae discussed below. Reared females,

mated if possible, were fed on 1:2 honey:water for at least a week before

being introduced singly to single hosts in clean containers, separately both naked

and in situ with foodplant. Providing they were fed adlibitum and protected

from extremes of temperature, most female Rogadinae lived in captivity for

about eight weeks of summer. The potential hosts offered were selected to

investigate both ecological and phylogenetic aspects of host range, as well as

to evaluate literature records. As far as possible the host(s) from which the

Shaw: Evolution of endoparasitism 309

species had been reared were also used, especially to investigate which host

stages were acceptable and suitable for successful parasitization. Although

subjective, evaluations of host acceptibility (of different instars as well as

different species) were surprisingly easy to make, almost all of the very many

female Rogadinae tested proving to be fairly decisive in behavior, and in close

agreement if conspecific.

RESULTS

Tribe EXOTHECINI

All species of Exothecini discussed have the following features in com-

mon: (a) destructive host-feeding is apparently obligatory, though light in

Rhysipolis, and often involves hosts too small for oviposition; (b) naked hosts,

as opposed to those in situ in plant material, are consistently and completely

ignored; (c) very many minutes, or even hours, are usually spent investigating

a host retreat for the presence of a host, and then in trying to sting it; (d)

the effects of the venoms on the hosts, as described below, are irrespective

of actual oviposition; (e) in all solitary species there is an ability to use the

larger hosts for female progeny; (f) egg hatch and larval development are rapid

(often only 4-8 days between oviposition and cocoon formation); (g) the larvae

feed entirely ectoparasitically; and (h) the winter is spent as a prepupa in the

cocoon.

(Genus GNAPTODON Haliday)

Although this genus is included in the Exothecini by Shenefelt (1975), I

fully concur with Achterberg (in prep.) that Gnaptodon should be excluded from

not only the Exothecini but even the Rogadinae as a whole. Gnaptodon species

are parasitoids of Nepticulidae (Lep.), and appear to be endoparasitic (although

this is yet to be thoroughly investigated).

1. Genus COLASTES Haliday (= Exothecus Wesmael)

Colastes braconius Haliday (type-species)

In Britain a widespread, abundant and regular parasitoid of a broad range

of leaf-mining insects, especially of the following groups: many Agromyzidae

(Dipt.) (not intensively sampled by me), most British species of which mine

plants in the field layer; Rnynchaenus (=Orchestes) fagi (L.) (Col-: Curculioni-

dae) on Fagus sylvatica; and all adequately sampled Phyllonorycter (Lep.:

Gracillaridae) species, arboreal and otherwise, except miners in bark or tough

evergreen leaves.

Less frequent as a parasitoid of various Nepticulidae, from which group

as a whole it has an abnormally male-biased sex-ratio (Shaw and Askew. 1976

and unpublished); Tischeriidae (Lep.) Tischeria ekebladella (Bjerkander) on

Quercus robur and T. marginea (Haworth) on Rubus fruticosus); and Mompha

raschkiella (L.) (Lep.: Momphidae) on Chamaenerion angustifolium.

Rarely (once each) from Eriocraniidae (Lep.) (Dyseriocrania sub-

purpurella (Haworth) on QO. robur); Heliozelidae (Lep.) (Heliozela betulae

310 Contrib. Amer. Ent. Inst., vol. 20, 1983

Stainton) on Betula pubescens and Antispila sp. on. Swida sanguinea); leaf-

mining Tenthredinidae (Hym.) (Fenusaulmi Sundewall on Ulmus and

Heterarthrus vagans (Fallén) on Alnus glutinosa); and Rhynchaenus species

other than fagi ['R. avellanae (Donovan) on Q. robur].

Most other groups of British leaf-miners, except for Elachistidae (Lep.),

have been well sampled (see in part Askew and Shaw 1974).

It may be significant that, among the Lepidoptera, none of the leaf-

mining groups of Dacnonypha or Monotrysia are important hosts, the specializa-

tion of C. braconius being at variance with the life-histories of these. With

the proviso that I have very incomplete information on the most frequently

used Diptera, the important hosts of C. braconius were seen to (a) complete

their development and pupate in a single mine; (b) inhabit blotch mines that

eventually develop some degree of depth (even if only to accommodate the

host cocoon, pupa, or puparium); and (c) do not isolate the pupation chamber

of R. fagi is a merely weakly isolated swelling in the leaf apex). Thus

species which change leaves, or pupate outside the mine, or strongly isolate

a small pupation chamber as a reinforced and sometimes deciduous disc or

sphere, are to a greater or lesser extent unsuitable for efficient exploitation

and outside the host range. C. braconius first emerges in April or May to

have several summer generations: to a large extent different host gruops are

attacked at different times of year.

C. braconius was examined experimentally as a parasitoid of both R.

fagi, and Phyllonorycter species onQ. robur and Betula. The host is most

often attacked as a post-feeding larva, but can be parasitized successfully also

during late feeding instars, or as a prepupa if only weakly cocooned. The fe-

male parasitoid probes the mine, apparently at random at least initially, re-

peatedly withdrawing and reinserting the ovipositor. Dead or otherwise inactive

hosts are rejected, but an active host is stung when located and the ovipositor

is withdrawn: a few seconds later the mine is again penetrated and a single

egg is laid in the mine, apparently placed at random, and typically well away

from the host. The venom received by the host causes an immediate paralysis,

from which a temporary and rather variable partial recovery is often made

within a few minutes. During the post-paralysis period (up to c. 1 day) the

host usually remains still, and never resumes feeding or cocoon construction,

although it is in some cases capable of considerable movement if disturbed.

Defecation takes place and the gut usually becomes fully vacated, but the po-

tential for activity quickly declines until, by the time the parasitoid egg

hatches, the host is usually apparently dead. The first instar parasitoid larva

has well-developed antennae and travels through the mine to the immobile host,

on which it feeds entirely externally throughout its development on all hosts

studied. [This refutes Beirne's (1946) contention, perpetrated by various sub-

sequent authors, that it is-endoparasitic]. Cocoon formation often occurs well

away from the host remains, usually at the extreme edge of the mine. Mobile

hosts are attacked irrespective of whether or not they are already endopara-

sitized but, as subactive hosts are usually rejected, superparasitism rarely occurs.

2. Genus PHANOMERIS Foerster

Phanomeris catenator (Halliday) [= ? dimidiatus (Nees) = ? abnormis

(Wesmael), type-species]

Shaw: Evolution of endoparasitism ott

In Britain a widespread and locally abundant parasitoid of the following

arboreal leaf-mining sawflies (Hym.: Tenthredinidae, Fenusini): Parna tenella

(Klug) on Tilia; Messa hortulana (Klug) on Populus nigra ; M. nana

(Klug) on Betula; Profenusapygmaea (Klug) on Quercus; and Fenusa ulmi

on Ulmus . Like other Fenusini, the hosts feed in shallow single mines and

undergo post-feeding ecdysis, usually before leaving the mine (or in F. ulmi

often immediately afterwards), to become a non-feeding eonymph which con-

structs a cocoon in the soil in which pupation occurs. P. catenator first

emerges in May and is plurivoltine. Some of its hosts (e.g., P. tenella ) are

available at only particular times of year, but it is fully synchronous with

others (e.g., P. ulmi).

In experiments P. catenator consistently refused to attack Heterarthrus

aceris (Kaltenbach) and H. vagans (both Hym.:Tenthredinidae, Heterarthrinae),

Or Scolioneura betuleti (Klug), Fenusa pusilla (Lepeletier), Py aohrnii

(Tischbein) and Fenella nigrita Westwood (all Fenusini). It was investi-

gated in culture only as a parasitoid of F. ulmi . Host mines nearing full

growth or less than half grown are consistently rejected (except for host-

feeding), as are all subactive hosts. Mines from about half grown are probed

repeatedly, initially at random (mines with their hosts removed were probed

for up to 10 minutes before being abandoned), when the surprisingly mobile

host usually wriggles away with ease. When eventually contacted, the host

is stung and paralyzed: the parasitoid withdraws its ovipositor and a few

seconds later again inserts it into the mine, laying an egg usually away from

the host. The host recovers from the paralysis within a few minutes and

usually returns to a feeding position at the edge of the mine but, although

movement of its mandibles can be seen, it does not resume feeding to any

appreciable extent. It remains moderately and voluntarily mobile for up to

c. | day, and always defecates until its gut is cleared, eventually withdrawing

from the edge of the mine and becoming immobile. The newly hatched para-

sitoid larva is similar to that of C.braconius and quickly locates the mori-

bund host. The cocoon is a tough, spindle-shaped single envelope structure

spun against both surfaces of the mine, generally near but not fully at the

mine boundary. Parasitized hosts in their period of renewed mobility are

sometimes superparasitized but it is extremely rare for more than one cocoon

to result (except in multi-host, confluenct mines). Mobile hosts harboring

endoparasitic Chalcidoidea are not discriminated against and, as is also seen

in the case of C. braconius, frequently both species develop successfully as

multiparasites.

Phanomeris laevis (Thomson)

In southern and central Britain, a local but sometimes abundant parasi-

toid of Heterarthrus aceris on Acer in midsummer, and subsequently both

H. vagans and Fenusadohrnii on Alnus in their autumn. The Heter-

arthrinae differ from the Fenusini in their pupation habits. The similar post-

feeding ecdysis to an eonymph is followed by its strong isolation in a small

reinforced disc within the mine, in which pupation later occurs. In H. aceris

the disc falls from the leaf but in H. vagans it does not. P. laevis first

emerges in early June and is plurivoltine (? bivoltine), using chiefly the uni-

voltine H. aceris in its first generation.

ol2 Contrib. Amer. Ent. Inst., vol. 20, 1983

In culture this species has been investigated only as a parasitoid of

H. aceris. The biology is essentially the same as that of P. catenator: the

host is most suitable from about half grown and host mines are rejected as

they approach full growth; hosts are stung and paralyzed at first but they re-

cover some activity, voiding the gut within a few hours but not resuming

feeding, and remaining largely quiescent until again becoming immobile after

c. 1-2 days; and the parasitoid egg is laid loose in the mine often some dis-

tance from the host, which is located by the first instar larva. The cocoon

is a broader, double envelope structure, often spun near the edge of the mine.

3. Genus XENARCHA Foerster

Xenarcha lustrator (Haliday) (type-species)

A locally common parasitoid of the leaf-mining sawfly Fenella nigrita

in southern England, on both Agrimonia eupatoria and Potentilla reptans.

X. lustrator first emerges in June/July and is plurivoltine (? bivoltine), being

well synchronized with its host.

In culture X. lustrator was offered F. nigrita mines on both its food

plants. Mines about half to three-quarters grown are accepted for oviposition,

those larger than this being consistently rejected. The mine-probing and host

location patterns are similar to those of Colastes and Phanomeris. The host

is temporarily and only partially paralyzed when stung, and the parasitoid lays

one or sometimes up to three (i.e., during a single visit but probably involving

separate insertions of the ovipositor) eggs loose in the mine. The initial

paralyzing effect of the venom is short-lived and the host resumes feeding

within c. 5 minutes, somewhat enlarging its mine during the tollowing 6-12

hours before withdrawing towards the center of the mine where it becomes

quiescent, though clearing the gut by defecation and being capable of movement

if disturbed for a further day or two. The egg(s) of X. lustrator are some-

what sticky and tend to collect on the host as it contacts them during its

period of temporarily recovered activity, so that the egg(s) are usually found

on the host by the time it has again become inactive. (This tendency, cer-

tainly strong in both species of Xenarcha, was noticed on a reduced scale in

Phanomeris, but seemed to be very slight or absent in Colastes. This prob-

ably reflects the relative activities ot the hosts stung by these parasitoids).

The egg(s) sometimes hatch before the inactive host is actually dead. The

cocoon is a stout, slightly curved, bluntly spindle-shaped structure spun near

the edge of the mine. Irrespective of the number of eggs laid, only one co-

coon resulted from each Fenella mine: an attempt was made to see if

Metallus species, also on low-growing Rosaceae but larger than Fenella, would

be attacked and perhaps permit gregarious development but, although one larva

ot M. gei Brischke in Geum urbanum was stung and suffered the same pattern

of vacillating activity, no eggs were laid.

Xenarcha, sp. indet.

An extremely local parasitoid of Fenusa pusilla on Betula and F.

dohrnii on Alnus glutinosa, found only (but in F. pusilla abundantly) at

respectively two sites in southern England and one site in north west England,

always virtually contined to mines close to the ground.

Shaw: Evolution of endoparasitism 313

In culture mines of Heterartherus vagans and F. dohrnii (both alnus

glutinosa), H. microcephalus (Klug) (on Salix aurita), H. nemoratus

Fallén), Profenusa thomsoni (Konow), Scolioneura betuleti and Messa nana

(all betula), Profenusa pygmaea (on Quercus ) and Fenella nigrita (on

Potentilla reptans) were all rejected; though some of the mines of F.

dohrnii, and Heterarthrus on both Alnus and Betula, were probed. Females

reared from both F. pusilla and F. dohrnii successfully attacked F.

pusilla. The biology is essentially as in x. lustrator, except that the host

is acceptable only when the mine is about half grown or under; the initial

paralysis of the host is even less complete and shorter-lived; the subsequent

feeding period by the host results in as much as 100% mine enlargement and

lasts for c. 2-3 days before the host rather abruptly ceases activity; and no

tendency to lay more than one egg per host was noted (although more than

one host is often parasitized in multi-host confluent mines). The adult

emerges in June/July and is long-lived: there seems to be only a partial sec-

ond generation.

4. Genus ONCOPHANES Foerster

Oncophanes laevigatus (Ratzeburg) [= lanceolator (Nees); it is probable that

laevigatus is also identical with the type species, minutus (Wesmael) ].

A widespread and, especially in the south of Britain, abundant gregarious

parasitoid of middle and late instar leaf-rolling Tortricidae and other small

lepidopterous caterpillars of similar habits. Reared from many arboreal tor-

tricids (unidentified species on Prunus, Crataegus, Sorbus, Malus, Betula and

Ulmus), and also Agonopterix nervosa (Haworth) (Oecophoridae) on Ulex

europaeus; and once each from the following Lepidoptera which are beyond

its normal host range but include evidence that hosts are also sought in the

field layer: Scrobipalpaacuminatella (Sircom) (Gelechiidae) on Cirsium

palustre, Prochoreutis sehestediana (Fabricius) (Choreutidae) on Scutel-

laria galericulata, and Phyllonorycter sp. on Salix aurita. The usual

hosts are mobile, in the sense of periodically constructing new retreats from

which feeding sorties may be made, and mostly pupate among the leaves of

their foodplant. O. laevigatus first emerges in May and is plurivoltine: it

probably attacks different host species at different times of year, as many

tortricids are univoltine.

In culture O. laevigatus has attacked all the species of Tortricidae in

rolled, folded or spun leaves that have been offered [unidentified species on

Populus alba, Hypericum perforatum and Malus sylvestris, and Epinotia

immundana (Fischer von Roéslerstamm) on Alnus glutinosa]. The host is stung

through the leaf, and is permanently paralyzed. The parasitoid then enters

the leaf-roll (etc.) and remains inside by the host, on which it host-feeds,

for up to | day before re-emerging: oviposition occurs towards or at the end

of this period. The eggs are slenderer and more curved than in the genera

already discussed, and are arranged more or less precisely along the (sub)

ventral aspect of intersegmental membranes, between several usually fairly

central adjacent segments of the host's body. The eggs are somewhat sticky,

but not firmly glued, and are easily removed. The host remains alive (for

up to 10 days in the absence of parasitoid eggs), pulsating slowly, but has no

gross mobility. The spindle-shaped papery cocoons are spun as an adpressed

314 Contrib. Amer. Ent. Inst., vol. 20, 1983

mass in the host retreat, usually flattened to the leaf surface(s). Brood sizes

of c. 4-15 are most usual, and the adults remain loosely associated with

their cocoons for a time after emerging, indicating that sib-mating may be

habitual.

5. Genus RHYSIPOLIS Foerster

Rhysipolis decorator (Haliday)

In Britain a widespread and common parasitoid of Caloptilia alchi-

miella (Scopoli) (Lep.: Gracillariidae) and C. robustella Jackh on Quercus in

autumn, also attacking C. betulicola (Hering) on Betula especially earlier

in the year. A very few male specimens have also been reared from confirmed

Phyllonorycter mines on Quercus and Betula in the course of very intensive

sampling (Askew and Shaw 1974, 1979; Shaw and Askey 1976), but these small

species are outside the host range. One female reared from the poorly sampled

Acrocercops brongniardella (Fabricius) (Gracillariidae) on Quercus may indi-

cate an important, if local, midsummer host. R. decorator is bivoltine, first

emerging at the end of May. Closely related and biologically similar forms

attacking (a) C. elongella (L.) on Alnus and probably also Cc. betulicola on

Betula, (b) C. rufipennella (Hiibner) on Acer pseudoplatanus (in the Nether-

lands and Switzerland, but not so far found in Britain), and (c) Parornix

(Gracillariidae) species on Prunus and perhaps Crataegus, are probably each

distinct species. The usual hosts of R. decorator make a single small mine

in their early instars, later feeding and moulting inside a series of two to three

conical folds, constructed usually on successive leaves; and the substantial co-

coon is spun on the lower surface of the leaf usually some distance away from

the final cone. The host is attacked in its final cone, rarely its penultimate

cone, as either a penultimate or more often final instar larva.

Experimental sleevings of R. decorator from C. alchimiella with

Phyllonorycter quercifoliella (Zeller) on Quercus robur led to neither para-

sitization nor host-feeding. Caloptilia syringella (Fabricius) in Syringa rolls

was similarly rejected. Adults from C. alchimiella readily parasitized c.

robustella in sleeves and C. betulicola in tubes, infrequently host-feeding.

The adult parasitoid patrols the cone, repeatedly making rapid thrusts with

its ovipositor through the leaf and into the cavity, by which means it even-

tually stings the host and withdraws its ovipositor. The venom causes tempor-

ary paralysis, and oviposition takes place from outside the cone. The elongate

and strongly curved egg is placed snugly and accurately along an intersegment-

al membrane, usually between abdominal segments. It is glued so strongly that

it cannot be detached without damage. Superparasitism is frequent - even

usual - both in the field and in culture, and appears to take place during a

single visit, but only one parasitoid cocoon per host ever results. Within about

an hour the host recovers full mobility and continues to feed; however, at the

end of the feeding period of that instar, whether it is nominally penultimate

or final, the host spins a cocoon inside its cone (in an experimental sleeve

of C. robustella a few of the parasitized hosts made cocoons outside their

cones, as is normal, but in all other cultures and in the wild the cocoon ap-

pears to be almost invariably constructed in the cone attacked by R. decora-

tor), inside which its development is arrested in prepupal proecdysis. The

parasitoid egg sometimes hatches before the host spins its cocoon; in either

Shaw: Evolution of endoparasitism ro ee.

case, the first instar larva does not fully emerge, its caudal half remaining

within the somewhat horny eggshell (which collapses only posteriorly) during

the early part of its feeding, and it only fully quits the eggshell in a later

instar after the host has become moribund in its cocoon. Despite its proxi-

mity to the host's body at all times, the head of the first instar larva bears

antennae comparable with those of other Exothecini. R. decorator makes its

own cocoon, in appearance very similar to that of a solitary Oncophanes, with-

in that of its host.

Rhysipolis hariolator (Haliday)

Widespread and locally common in Britain as a parasitoid of Parornix

betulae (Stainton) on Betula and P. devoniella (Stainton) on Corylus avel-

lana, Previous records from Phyllonorycter species on Corylus (Shaw and

Askew 1976) may have been in error: the mine and fold of P. devoniella

each closely resemble mines of this genus. The host larva makes a tentiform

mine which it leaves in its final instar to make one or more folds at this or

another leaf edge, in which it completes its feeding. Pupation takes place

in a small tight fold at the edge of a different leaf, either of the foodplant

or beneath it. Both of these hosts are bivoltine in southern Britain where

R. hariolator is also bivoltine and well synchronized, first emerging in May/

June.

The host is attacked as a final instar larva, perhaps exclusively so, in

both the mine and the fold: hosts attacked in the mine proceed to a folding

stage. Egg placement and the tendency to superparasitism are as described

for R. decorator, and similarly the host resumes feeding after initial paralysis

only to die in its cocoon which is, however, usually spun in its normal situa-

tion away from the last fold.

Rhysipolis ?meditator (Haliday) (type-species)

Widespread and abundant in Britain as a parasitoid of Mompha rasch-

kiella (Zeller) (Lep.: Momphidae) mining Chamaenerion angustifolium; a

single rearing from M. terminella (Humphreys and Westwood) mining Circaea

lutetiana suggests that a wider range of leaf-mining Momphidae may be used,

but only M. raschkiella has been adequately sampled. The latter makes

blotch mines and is able to change leaves, probably usually doing so at least

once or twice before it is fully fed. It pupates in a cocoon spun in debris

or a tightly folded leaf near the ground, and is largely bivoltine. R.

?meditator is well synchronized and similarly chiefly bivoltine, first emerging

-in June.

The host is attacked as a final instar larva, possibly exclusively so. Egg

placement and the tendency towards superparasitism are as described for

R. decorator. The host dies in prepupal arrest in its cocoon, which is almost

invariably spun in its usual situation, inside which the parasitoid makes its own

cocoon. Eggs of the parasitoid have not been found to hatch before host co-

coon formation and preliminary observations suggest that, within the attacked

instar, feeding by parasitized larvae (which is certainly resumed) may be cur-

tailed compared with unparasitized individuals. Although any such tendency

was not noted in R. decorator or R. hariolator, this requires further investi-

gation in all three species.

316 Contrib. Amer. Ent. Inst., vol. 20, 1983

Rhysipolis, spp. indet.

Two as yet unidentified, and biologically uninvestigated, species have been

reared from cocoons of (a) Phyllonorycter emberizaepenella (Bouche) mining

Lonicera periclymenum (Netherlands, not so far in Britain) and (b) Phyl-

lonorycter ? schreberella (Fabricius) mining Ulmus (Switzerland, not so far

in Britain). It is of considerable significance that these two hosts differ from

most Phyllonorycter species in constructing strong cocoons in the mine (which

are further protected by being only feebly connected to the leaf tissue), no

doubt aiding overwintering in the face of the abnormal susceptibility of these

leaves to fungal decomposition.

Tribe ROGADINI

1. Genus CLINOCENTRUS Haliday

Clinocentrus gracilipes (Thomson)

In Britain a widespread and abundant parasitoid of all sampled Choreuti-

dae (Lep.) in low vegetation: Prochoreutis sehestediana and P. myllerana

(Fabricius) on Scutellaria galericulata, and Anthophila fabriciana (L.) on

Urtica. Prochoreutis species have two summer generations, at least in south-

ern Britain, larvae appearing in June/July and again in July/August. The larval

stage almost certainly does not overwinter, though the life-histories of these

two moths are incompletely known. A. fabriciana overwinters as an early

larval stage (Shaw 198la) and is fully grown by June after four larval instars:

subsequently there are one to two further summer generations. Both

Prochoreutis and Anthophila spin webs on leaves of their foodplants, causing

them to form boat-like concavities or open pods, in which the larva rests and

feeds in semi-concealment. About two or three leaf changes are usual during

the larval life; and subsequently a dense, silken double cocoon is constructed

amongst debris on the ground or in a more or less folded leaf, usually below

and well-distanced from the feeding site. C.gracilipes is largely bivoltine

and overwinters as a prepupa inside the "mummified" host prepupa in its co-

coon, from which it first emerges in mid to late June. Synchronization of

C. gracilipes with its ubiquitous host, A. fabriciana, on which it is usually

locally entirely dependent, is very poor as the overwintering host generation

is almost completely missed. As a parasitoid of Prochoreutis it is far better

synchronized, but Prochoreutis species are extremely local, occurring chiefly

in fens, and support only a minute fraction of the British population of

C. gracilipes.

In culture C. gracilipes readily and successfully attacks Choreutis

pariana (Clerck) (Choreutidae), which feeds on rosaceous trees in a similar

way, but it has never been reared from extensive collections of this arboreal

host (e.g., Shaw, in press and unpublished) even at sites where it was causing

heavy mortality to A. fabriciana on Urtica beneath the trees. Various

tortricid and yponomeutid larvae of apparently similar habits were consistently

rejected. A. fabriciana was used for detailed experimental rearings, using

parasitoids reared from both Prochoreutis and Anthophila. Third and fourth

(= final) instar larvae in their webs are accepted, sometimes even if subactive

in proecdysis, but naked hosts are consistently ignored. The adult probes the

Shaw: Evolution of endoparasitism St7

host web with its ovipositor apparently at random (and indeed it makes lengthy

searches of recently vacated webs) and from many angles, periodically making

sharp stabbing thrusts, and rapidly withdraws the ovipositor after eventually

stinging the host. The venom rapidly causes temporary paralysis, during which

the adult parasitoid relocates the host and oviposits through the web and into

its body. Host-feeding has never been seen to take place, and nor is there

a tendency towards superparasitism at a single visit. The egg is laid in a

transverse position immediately beneath the epidermis, to which it adheres,

and is clearly visible externally. Except that the egg occupies a central posi-

tion in a body segment and is placed below the skin, egg placement is remark-

ably similar to that of Rhysipolis, and the following effects of the venom

(Shaw 198lb) are identical: after temporary paralysis the host recovers mobil-

ity and feeding activity (seemingly undiminished, but precise data lacking) and,

at the end of the feeding period of whichever instar is attacked, it is switched

to a prepupal state in which it exhibits full cocoon-making behavior before

eventually dying as a pharate pupa in developmental arrest, the body contents

having undergone considerable histolysis. The parasitoid egg often hatches be-

fore the host becomes arrested but the bulk of larval development takes place

afterwards, and the fully grown parasitoid larva forms its cocoon within the

skin of the host. This "mummy" is a dark, hardened and uncontracted struc-

ture comprising the host larval skin fairly substantially lined with silk, and the

adult parasitoid emerges from it by biting a hole just behind the hosts's head

but not at any particular dorsoventral orientation. At the time of mummifica-

tion a small quantity of fluid sometimes escapes as seepage near the caudal

end of the host, but the mummy does not become glued down in any way.

€, gracilipes shares with all investigated Exothecini the following fea-

tures: (a) it attacks only concealed or semi-concealed late instar hosts in

situ; (b) it has a lethal venom controlling subsequent host behavior and develop-

ment, which is applied as a separate action before oviposition; (c) it has rapid

larval development upon a moribund and physiologically static host; (d) it is

able to respond to the size of the host attacked by laying female eggs pre-

dominently in the larger host individuals and male eggs mainly in the smaller

ones; (e) it invests a considerable period of time in each oviposition sequence;

and (f) it overwinters as a prepupa in its cocoon.

Remarkably little is known of the host associations of other species of

Clinocentrus, but probably "microlepidoptera" living in or under webs or in

other spun retreats, but perhaps pupating elsewhere, will be the normal hosts.

I have seen reared specimens of C. excubitor Haliday from Tortricidae and

C. vestigator Haliday from Yypsolopha vittella (L.) (Yponomeutidae), and

it is possibly significant that leaf-mining occurs in both of these lepidopterous

families (admittedly mainly in early instars in the case of Tortricidae), and

also in some (non-British) Choreutidae.

2. Genus PELECYSTOMA Wesmael

No species of Pelecystoma has been reared by me, but details given by

Smith et al. (1955) for the Nearctic P. harrisinae (Ashmead) suggest that

it occupies a biologically intermediate position between the otherwise widely

spaced genera Clinocentrus and Aleiodes/Rogas. Rearing records given by

Shenefelt (1975), including those for the type species P. luteum (Nees) and

318 Contrib. Amer. Ent. Inst., vol. 20, 1983

the gregarious (see Watanabe 1962) P. tricolor Wesmael from the Palearctic,

suggest that Pelecystoma attacks fully exposed larvae belonging to "primitive"

families of Lepidoptera (Limacodidae and Zygaenidae), which might suggest

that this genus arose through an early response to the appearance of the naked

feeding habit in Lepidoptera. |

3, 4. Genera ALEIODES Wesmael and ROGAS Nees

As discussed by Shenefelt (1969), the correct application of generic

names in this group is problematical: here they are applied sensu Shenefelt

(1975) and, as no significant overall difference between the biologies of

Aleiodes and Rogas has been evident from the fairly numerous species that

I have studied, the two genera are treated together. However, there is un-

doubtedly a broader total host spectrum and more apomorphy in terms of life-

history adaptations (in particular regarding overwintering) seen in Aleiodes

than in Rogas. The latter genus seems to be chiefly univoltine and associated

with Noctuidae.

In general, both genera (a) attack early instars of exposed "macrolepi-

doptera" (Hesperoidea, Papilionoidea, Bombycoidea, Geometroidea, Sphingoidea,

notodontoidea and Noctuoidea in Britain); (b) exhibit various degrees of host

specificity, voltinism and overwintering habits; (c) attack hosts away from

foodplant substrates as readily as on them; (d) employ venoms that cause short-

lived temporary paralysis as a handling device facilitating subsequent oviposi-—

tion, but having no other detectable physiological effect on the host; (e)

oviposit loosely into the host haemocoel as a separate action distinct from in-~

jecting venom; (f) kill the host in a middle instar, i.e., parasitized, making

a somewhat contracted mummy; and (h) host-feed non-destructively and only

when deprived of other nutritive moisture. Some illustrations of, and excep-

tions to, these generalizations are given below:

(a) In addition, A. bicolor (Spinola) is a regular parasitoid of Zygaena

filipendulae (L.) (Zygaenidae) in which it overwinters but, as it also attacks

Polyommatus icarus (Rottemburg) (Lycaenidae) on the same plant (Lotus) in

summer, the use of Zygaena may be a secondary adaptation.

In experiments with A. ?circumscriptus (Nees) and Orthosia stabilis

(Denis and Schiffermuller) (Noctuidae) it was found that mid-first, mid-

second and early third instar hosts were approached with similar enthusiasm,

but that first instar hosts were attacked only with difficulty and a significant-

ly higher proportion of them did not survive the oviposition sequence itself.

Development time to host mummification was shortest in third instar hosts

(0-1 months) and longest by far in first instars (2-3 months). Mean handling

time (i.e., the whole oviposition sequence) was shorter in second instars than

in either first or third instars but the range was too great to assess signifi-

cance. Encapsulation was significantly higher in third instar hosts than in

firsts and seconds. The sex-ratio of adults reared from the three instars did

not differ significantly although rather larger individuals resulted from the

later instar hosts. Less detailed experiments with A. geniculator (Nees)

parasitizing Orgyiaantigua (L.) (Lymantriidae) and Spilosoma (Arctiidae)

species indicated similar trends, as did experiments involving R. rugulosus

(Nees) as a parasitoid of Acronicta rumicis (L.) (Noctuidae).

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(b) Life-history data of some species for which I have most experience

are summarized in Table 1. The habit of using host species in which the

parasitoid is able to overwinter as an early instar larva in a living caterpillar

is best developed in Aleiodes, although it is also seen in certain Rogas species

not treated in detail here. Overwintering prepupae in mummies suffer high

rates of predation and parasitism, and it is probably significant that the

species listed in Table | that overwinter as mummies are on the whole far

less abundant than those that do not.

It was found that, as in many other parasitoids, there is in almost all

cases a very strongly defined division into arboreal species, and those on low

vegetation. Thus the host ranges comprise groups of hosts in one or other

of these partitions of the environment, closely related species from other

niches being completely free of the parasitoid in nature. These "hosts" are,

however, often attacked if offered in culture, especially after repeated or pro-

longed contact, but parasitoid survival is extremely poor. This probably re=-

flects a lack of the regular contact that might lead to adaptation, but it also

shows that if a physically acceptable potential host newly arrives in the para-

sitoid's searching environment it will be attacked and, if this happens frequent-

ly enough, it will presumably generate sufficient selection pressure to result

in compatibility and hence an adoption of that host into the host range. Al-

though rather limited in this context of host range extension, this does sug-

gest ways in which resource abundance might indeed trigger behavioral res-

ponses relevant to the hypotehsis that the availability of behaviorally novel host

groups has been an important evolutionary influence on the Rogadinae as a

whole.

Noteworthy details for particular Aleiodes species are: (i) the excep-

tional and rigid seasonal alternation between arboreal and field layer hosts

by A. ?circumscriptus, probably first arising as a response to seasonality in

resource abundance; (ii) the thelytoky, probably associated with extreme host

specificity, of A. pallidator (Thunberg); and (iii) the overwintering as an

adult by A. testaceus (Spinola), enabling it to attack the very early spring

larvae of arboreal geometrids soon after they hatch from their overwintering

eggs.

traces of various kinds (feces, silk, chewed leaves), but hosts are often first

found visually (notably in A. geniculator and R. rugulosus) or by random

antennal contact, and all species tested accepted isolated hosts freshly placed

in glass tubes. Direct antennation of the host, and frequently also contact

with the fore and/or mid tarsi, is important in the host acceptance process.

The paralyzing sting (but see below) is brief, after which the ovipositor is with-

drawn, and both the anterior legs and the antennae are used to manipulate

and hold the host in position during the rest of the oviposition sequence. The

adult parasitoid waits (c. 40-60 sec) until after paralysis has taken effect

(c. 5-15 sec) before again inserting the ovipositor, usually only once, for

oviposition (c. 30-80 sec). After oviposition the parasitoid usually remains

standing over the host (c. 20-100 sec), sometimes antennating it; this might

possibly be interpreted as guarding the host or keeping it associated with its

substrate until the effects of paralysis start to wear off, although the adult

Shaw: Evolution of endoparasitism 321

parasitoid had usually left the host rather well before any such recovery

(c. 2-5 min) was evident to me.

(d) All species studied, except for A. heterogaster Wesmael and A.

praetor (Reinhard), use a venom to induce temporary paralysis. The lack of

paralysis by these two species is probably an adaptation allowing their often

monophagous arboreal hosts to continue to cling on to the foodplant while

oviposition takes place. All other species studied (including those that are

arboreal) insert the ovipositor twice - once to inject venom, then again to

oviposit - and, by switching host individuals at the appropriate time in the

oviposition sequence, oviposition into hosts that had not received venom could

be induced: in these cases (A. ?circumscriptus, A. ?nigriceps Wesmael,

A. geniculator, and R. rugulosus) parasitoid development was absolutely nor-

mal with no evident increase in encapsulation levels. Similarly, if stung hosts

were removed before oviposition took place their development was completely

normal and lead to healthy pupae whose formation lagged behind controls by

only a few hours, probably representing no more than the full recovery time

following paralysis [this was confirmed on a small scale for many species of

both genera; in detail for A. ?:circumscriptus, A. modestus (Reinhard) and

R. grandis (Giraud) ].

(e) Egg placement appears to be at random into the host's haemocoel

in all species studied. However, the process of actual oviposition is prolonged

enough to suggest that some tissue discrimination, at least, may occur: it

is, of course, also possible that this prolongation might indicate that some fur-

ther glandular secretion (e.g., for egg protection) is injected along with the

egg at this, rather than the previous, insertion of the ovipositor. If this were

to be the case, however, any gross controlling effect by the venom on host

development could still be ruled out, as hosts that successfully encapsulated

their parasitoids have all developed normally.

Provided they are active, parasitized hosts are accepted for oviposition

without discrimination: subactive hosts, including those recently paralyzed, are

avoided but otherwise post-oviposition displacement activity by the female

parasitoid appears to be the only behavior tending to reduce superparasitism.

No positive tendency to superparasitize a host at a single visit was seen except

possibly in A. heterogaster.

(f) The presence of a parasitoid larva in the host retards its growth

and prolongs the period spent in each instar, so that roughly half grown para-

‘sitized hosts are often mummified at about the time that unparasitized siblings

pupate. The host instar killed, which within a particular association depends

to some extent on the instar attacked, does not differ uniformly in the two

genera, slightly abnormal lateness in some Rogas species (e.g., R. dimidi-

atus (Spinola) and R. grandis) being offset by extreme lateness in Aleiodes

modestus which kills its very small host species as prepupae. Mummy struc-

ture is rather variable but usually involves some contraction of host segments

at either the head or caudal end, or both, and the mummy is usually glued

onto its substrate via a ventral opening in the thoracic region: this opening

is made even by some species (e.g., A. modestus, A. pallidator) in which the

mummy does not become glued down. A. heterogaster is a notable anomaly

as it produces an inflated structure that is partly an air sac, from only the

caudal half of the host. This is possibly an adaptation to the wetland

322 Contrib. Amer. Ent. Inst., vol. 20, 1983

habitats in which A. heterogaster often occurs, as the mummy is not glued

down to the substrate, and floats in water. The quantity of silk spun as a

lining in the mummy is variable, probably reflecting little more than the

length of time spent in the mummy, and shows no consistent generic differ-

ence. The most silk-free mummies, however, are seen in Aleiodes.

(g) Emergence from the caudal end of the mummy is the rule for all

the Aleiodes and Rogas species studied, individual exceptions being rare in both

genera. This contrasts with Clinocentrus in which all mummies | have seen

(Cc. excubitor as well as C. gracilipes) have emerged from the head end.

(h) Host-feeding is very rarely seen, and is always non-destructive. It

usually involves rather large host individuals, after oviposition. Some species

have also fed occasionally on the host's oral secretions that often appear as

it is paralyzed. In general, however, honey: water is adequate for egg matura-

tion and well-fed individuals do not host-feed.

5. Genus PETALODES Wesmael

Petalodes unicolor Wesmael (type-species)

A widespread, but extremely local, species in Britain, reared from lar-

vae of the following Lepidoptera: Hydriomena furcata (Thunberg) (Geometri-

dae) on Salix and Corylus avellana, and Clostera pigra (Hufnagel) (Noto-

dontidae) on Salix. As larvae both hosts rest and feed beteer spun leaves:

H. furcata is univoltine and overwinters as an egg, being available as a young

larva early in spring; while Cc. pigra is more variable in its time of appear-

ance but feeds later in the summer and overwinters as a pupa. P. unicolor

is crepuscular and bivoltine, overwintering as an adult to oviposit in early

spring (an adaptation also seen in A. testaceus ) and appearing again in July

and then late August/September (before hibernation). The host range is prob-

ably a good deal wider than that noted above: certainly C. pigra appeared

to be absent from two sites at which H. furcata supported P. unicolor imply-

ing that different summer hosts would be involved. Undoubtedly all hosts, how-

ever, would feed in spun leaf packages or similar structures, and would probab-

ly be arboreal.

P. unicolor has been reared for several generations as a culture alter-

nating between H. furcata and C. pigra. The following notes apply to para-

sitism of the latter host. No interest whatsoever was shown for naked hosts,

but the host retreat (including feces and silk) elicited excitement whether or

not it was tenanted. After an external examination of the host retreat with

its short antennae, the parasitoid thrusts the whole of its abdomen (which in

the female sex is laterally compressed, blade-like and considerably extensile)

through the silk or into the leaf package, twisting it up to 90° in pursuit of

the host if it is between leaves or in narrow runs. When contacted, the host

is stung: temporary paralysis is rapid and the parasitoid quickly relocates the

host, again using its abdomen alone, and reinserts its ovipositor. Oviposition

itself takes about 90-250 sec, longer than in Aleiodes/Rogas, and the egg

is placed in the haemocoel but fastened by its narrow end to abdominal muscle

tissue. Unfortunately the effect of the venom alone has not been properly

investigated, but nothing has suggested that it differs from that of Aleiodes/

Rogas. As in a few species of Aleiodes, the mummy is not glued down

Shaw: Evolution of endoparasitism 323

although seepage occurs through a ventral thoracic opening, and it contains

virtually no silk. Emergence is as in Aleiodes/Rogas. Although biologically

rather close to Aleiodes/Rogas, P.unicolor differs in attacking only con-

cealed hosts, and also in its ill-understood egg placement behavior. Its aber-

rant morphology is readily understood in terms of the hosts attacked, and it

is of interest that Petaloides,probably having evolved from an Aleiodes -like

ancestor using naked hosts, retains a fairly short ovipositor in its specializa=

tion on concealed hosts.

DISCUSSION

The above account is limited to the biology of some British Rogadinae

and omits not only morphological data but also any information at all on very

many genera (and even tribes) of World Rogadinae. It cannot, therefore, be

seen as the basis for a rigorous phylogenetic study. The present hypothesis

is a great deal more simple: that the biological adaptations described above,

or very similar trends, led to particular conditions that provoked a transition

from ectoparasitism to endoparasitism in cettain Rogadinae. The emphasis

is on the kinds of interaction between host biology and parasitoid biology that

actually occur, and it is not intended to imply that the species studied are

themselves close to any sort of transition: extant species represent meta-

stable states of biological efficiency and may be far distant from one another

in evolutionary terms, being survivors of processes that have perhaps involved

powerful selection pressures, and much extinction of transient forms in the

face of competitive inefficiency or some other effective resource contraction.

This is bound to be the case if behavioral aberrations, at first intrinsically in-

efficient but highly compensated by the advent in a changing environment of

large and initially underexploited resources, precede physiological and morpho-

logical adaptations in the evolutionary process. Although direct evidence is

admittedly lacking, there are reasonable grounds to suppose that there has been

some sort of progressive evolution in the Rogadinae such that, for example,

the extant genera Colastes, Rhysipolis, Clinocentrus and Aleiodes, which

utilize progressively more "advanced" host groups, can be seen as illustrating

a sequence of development in the exploitation of new opportunities. However,

our knowledge of how the relevant host groups coevolved with their own re-

sources, or how this incompletely understood host phylogeny has been distrib-

uted in terms of space, time and relative abundance, is so hazy that it is im-

possible to do more than merely speculate on tendencies. Thus, while the

following summary (written in somewhat teleological terms for the sake of

brevity) indicates how this may have happened as a result of exploiting hosts

efficiently, and suggests some particular specializations that may have been

important, it must be taken in the spirit of a speculative suggestion rather

than as wild dogmatic claim.

The genus Colastes attacks the least "advanced" of the lepidopterous

hosts and in C. braconius contains the species with the broadest host range

of any of the Rogadinae studied, although this species has a long ovipositor.

and clearly specializes on fully grown hosts that live and pupate in their single

mines (a habit that seems to encourage mine depth for protective isolation

of the pupal stage). If the first instar parasitoid larva is mobile, precise egg

placement is unimportant in a closed mine which is not about to be vacated

324 Contrib. Amer. Ent. Inst., vol. 20, 1983

by the host, and possibly an ancestral habit would have been simply to ovi-

posit into the mine; however, better control is gained if the host is stung and

immobilized, thereby preventing it from taking advantage of the maneuver-

ability afforded by a deep mine to damage the ovipositor or the egg, and also

preventing the host's subsequent pupation and hardening of its cuticle. The

host's temporary recovery from paralysis, sufficient to defecate, is a further

refinement, and its slow as opposed to outright death allows the young para-

sitoid to feed on fresh rather than putrid food.

The genera Phanomeris and Xenarcha appear to be specialized and prob-

ably rather recent responses to sawfly miners, which moult at the time of

either leaving their flat mines to pupate elsewhere or else isolating themselves

for pupation in small discs in the mine. The venom used by Phanomeris pre-

vents the host from accomplishing this vital moult, thereby keeping it in its

mine, and this leaves egg placement unimportant. However, there is some se-

lection towards mines that are not fully grown, perhaps because the surprisingly

agile hosts escape for longer in full-sized mines, which will often be already

vacated in any case, and considerable time is invested in searching for a host

within each mine visited. In Xenarcha this is more clearly seen, and the venom

permits the host to recover considerable feeding activity and to grow in size,

thereby providing a larger food resource, while retaining the advantages of

attacking mines of a size most likely to contain relatively easily locatable

hosts. The evident stickiness of the loosely laid egg is perhaps one way of

ensuring that the first instar larva easily finds its food; mere oviposition pre-

cision being a useless strategy as the host subsequently moves away.

Oncophanes utilizes larger hosts than the other Exothecini studied and |

the tendency towards multiple oviposition is here successfully developed. Al-

though some mine when young, when at the stage attacked its tortricid (etc.)

hosts live in leaf-rolls (etc.) which are often partially open, and repeatedly

spin fresh retreats for themselves, eventually pupating in situ. A paralyzing

venom prevents host ecdysis or movement and ensures that parasitoid eggs are

not sloughed at ecdysis or simply left behind; and eggs placed ventrally along

host intersegmental membranes may be better hidden from small predatory

invaders of the leaf-roll, and also help to ensure that the first instar para-

sitoid larvae easily find the host and do not wander right out of the roll. The

tendency to use venoms that delay host death, providing a less putrid or even

living food resource, is expressed in Oncophanes by use of a venom causing

an even, long-term paralysis. (Gregarious development is known in some genera

of endoparasitic Rogadini not studied by me, and it is possible that develop-

ments of the trend seen in Oncophanes ~- i.e., gregarious ectoparasitism of

large hosts, involving precise egg placement and a controlling venom - may

have preceded a transition to endoparasitism. The genera Rhysipolis,

Oncophanes and Clinocentrus are morphologically close and, although the

biologies of the studies species of Rhysipolis and Clinocentrus seem closest,

there are vague literature records of gregarious development in some

Clinocentrus species. There is, of course, no firm evidence that all endo-

parasitic Rogadinae, or even all species of the poorly studied genus Clino-

centrus as accepted, comprise monophyletic groups. The view that the

Rogadini arose from within the Exothecini leaves the latter as a paraphyletic

group, irrespective of the possibility of the Rogadini being polyphyletic).

Shaw: Evolution of endoparasitism oeo

In Rhysipolis the primary specialization has been to take advantage of

leaf-mining larval hosts which pupate in a strong cocoon capable of providing

protection. By adapting to this situation, species of Rhysipolis have also been

able to utilize hosts of high mobility - i.e., those which frequently change

mines, or subsequently make leaf folds, independently of ecdysis - and its

hosts may pupate completely away from their feeding site. The egg is firmly

glued to the host integument, its alignment between body segments partly

concealing it in a position that not only affords it protection as the host is

exposed between mines or folds, but also prevents it from being scraped away

against the small hole through which some of the hosts quit their feeding

sites. The venom has several vital effects: firstly by causing temporary

paralysis it enables eggs to be placed and attached with precision; secondly

it permits the host to recover and to hide and feed until the end of its instar

but then arrests it before ecdysis, thereby ensuring that the egg is not lost;

and, most importantly, thirdly it switches the host to a prepupal phase so that,

whichever host instar is attacked, cocoon formation still takes place. The

parasitoid larva consumes the host in this arrested and protected state before

spinning its own cocoon inside that of the host.

The kind of precisely predictable, and physiologically weakened, state of

the arrested prepupal host of Rhysipolis is an important and provocative pre-

cursor to the advent of endoparasitism, with the added immediate advantage

of better egg concealment and protection, and also possibly allowing host-

derived nutrients to enter the egg through a hydropic chorion (unlike Exo-~

thecini, which lay well-yolked eggs, Clinocentrus does not host-feed).

Clinocentrus uses a venom that has precisely the same effect as that of

Rhysipolis, producing the same physiologically unchallenging and unvarying en-

vironment for its endoparasitic larva, and Clinocentrus even orientates the

egg in a similar way that now seems to serve no clear purpose (though the

behavior of the first instar larva is not known). The peripheral position of

the egg, however, may be important in terms of oxygen availability as the host

becomes physiologically inactive.

It is worth stressing here that there is little evolutionary opportunity for

endoparasitism of tenthredinid sawflies to have arisen in the Rogadinae in the

way that seems to have happened in association with lepidopterous hosts, be-

cause leaf-mining sawfly larvae undergo ecdysis immediately before cocoon

construction (and usually before leaving the feeding site) which is always un-

dertaken by a specialized non-feeding instar. Any venom, therefore, that

switched a sawfly larva to a prepupal state would provoke an ecdysis before

cocoon construction, which could only be survived by specialized (e.g.,

stalked and anchored, as in Tryphoninae) eggs of a kind not seen in Rogadinae.

It might, however, be even more relevant that the leaf-mining habit is evident-

ly a fairly recent apomorphous condition in sawflies, whereas it has much

deeper roots in the Lepidoptera in which it is mostly found in relatively

plesiomorphous groups. Similarly, any possible importance of dipterous hosts

in the evoluation of the Rogadinae can be dismissed: Agromyzidae are in-

cluded in the wide host range of C. braconius, but clearly leaf-mining is an

extreme apomorphy in the Diptera, and the Agromyzidae a very recent family

(Rohdendorf 1974: 305). In any case, the biology of Cyclorrhapha seems un-

conducive to extreme venom and egg placement specializations by ectoparasites.

Leaf-mining in Coleoptera is also a generally apomorphous habit from which

326 Contrib... Amer..eat. Inst, vol.20, 1983

little further specialized behavior has developed, although some mining species

move from one leaf to another and almost all pupate in the ground (Crowson

1981: 593). Indeed, as the behavior at metamorphosis of Coleoptera is broadly

suitable, it would not be surprising to find some Rogadinae, perhaps related

to Phanomeris or Xenarcha, specifically associated with phytophagous beetles -

possibly even as endoparasites. However, the view that endoparasitism in the

Rogadinae arose from an originally ectoparasitic stock attacking leaf -—miners,

and developed to the level seen in Aleiodes/Rogas in the course of adaptations

to new opportunities as their host group as a whole evolved in behavior, can

be applied only to lepidopterous hosts.

Adaptation to an endoparasitic way of life by Clinocentrus has depended

heavily on the venom system of its ectoparasitic precursors, but endoparasitism

was presumably also encouraged by the availability and exploitation of hosts

which feed in less concealed sites than those of the ectoparasites, as the pro-

tection of the egg, for example against desiccation, would be partly ensured

by the host's integument. Once established, however, endoparasitism raises

very different demands from ectoparasitism if it is to be adapted successfully

to fully exposed hosts. The ectoparasites tend to attack concealed, late in-

star hosts, and their protected progeny have high survival rates. Superparasi-

tism is inherently low because of long handling times, low fecundity associated

with large anhydropic eggs, and an ability to reject subactive hosts. On the

other hand, endoparasites of fully mobile hosts would not so easily be able

to avoid superparasitism and, furthermore, the first parasitoid larva present

would soon come to control the host's body well enough to destroy any subse-

quent eggs. As hosts that feed exposed became available and were adapted

to, they may well have attracted high levels of superparasitism, because of

much reduced handling times and the higher fecundities associated with this

and the scope for smaller, hydropic eggs. On this basis, natural selection would

favor endoparasitism of the host earlier in its life, as the first endoparasite

present would be at a competitive advantage, but for this to be taken very

far the host would subsequently have to be able to grow and moult so that

a sufficient food resource was eventually present. The controlling effect of

a venom that prevented host ecdysis would no longer be advantageous, but its

absence would provoke a further and power ful selection towards attacking

younger hosts with less copious physiological defenses against foreign bodies,

and also to specialize by evolving specific physiological tolerance towards nar-

row host ranges. (The striking difference between firstly Clinocentrus

gracilipes and secondly apparently all Aleiodes and Rogas species in their

abilities to develop in experimental hosts outside the host range - i.e., in this

case species closely related to the natural hosts but living in a different en-

vironment in which the parasitoid does not usually search -~ is an illustration

of this trend). The evolutionary distance between these two suits of biological

characters, befitting successful ectoparasitism and the first establishment of

endoparasitism on the one hand and successful endoparasitism of exposed hosts

on the other, thus involves various selective influences made particularly harsh

by being imposed in concert. This may account for the apparently large bio-

logical gap, perhaps partly filled by relict genera such as Pelecystoma, between

Clinocentrus and the again successful genera Aleiodes and Rogas, by which

the transition has been fully accomplished. In the latter two genera a throw-

back to an ectoparasitic ancestry in which a far greater role was played by

the venom, now only a temporary handling device which is still useful under

Shaw: Evolution of endoparasitism S27

most (but not all) circumstances, is perhaps seen in the separate insertions

of the ovipositor for paralysis and for subsequent oviposition. Killing the host

before it is fully grown may also be indicative that Aleiodes and Rogas suc-

ceeded chiefly by a fairly rapid progression to attacking the host very early:

some of the (unstudied) gregarious endoparasitic Rogadini, briefly mentioned

above in connection with Oncophanes, might, of course, turn out to be merely

extensions of the Aleiodes/Rogas form of development, taking advantage of

this extra host capacity.

ACKNOWLEDGMENTS

Part of the field and experimental work was carried out at the University

of Manchester supported by a grant (GR/3/864) from the Natural Environment

Research Council, and continued under the tenure of a University Research

Fellowship at the University of Reading. I am grateful to Dr. W. Junk, Pub-

lishers, for kind permission to reproduce parts of the discussion of host range

that first appeared in Series entomologica vol. 18, and to Drs. G. Bryan and

S. E. Whitebread for sending me Rhysipolis species reared from Phyl-

lonorycter mines in respectively The Netherlands and Switzerland. To Dr.

R. R. Askew I owe the greatest debt, for his interest and encouragement, and

for the opportunity he gave me to start on this work.

REFERENCES

Achterberg, C. van. 1976. A preliminary key to the subfamilies of the

Braconidae (Hymenoptera). Tijdschr. Ent. 119: 33-78.

Askew, R. R. and Shaw, M. R. 1974. An account of the Chalcidoidea

(Hymenoptera) parasitizing leaf-mining insects of deciduous trees in

Britain. Biol. J. Linn. Soc. 6: 289-335.

Askew, R. R. and Shaw, M. R. 1979. Mortality factors affecting the leaf-

mining stages of Phyllonorycter (Lepidoptera: Gracillariidae) on oak and

birch. 2. Biology of the parasite species. Zool. J. Linn. Soc. 67: 5l-

64.

Beirne, B. P. 1946. Notes on the biology of some hymenopterous parasites

of the beech weevil (Rhynchaenus fagi L.) (Col.). Proc. Roy. Ent. Soc.

London (A) 21: 7-Il.

Crowson, R. S. 1981. The biology of the Coleoptera. Academic Press.

Rohdendorf, B. 1974. The historical development of Diptera. Univ. of Alberta

Press.

Shaw, M. R. 198la. Overwintering of Anthophila fabriciana (L.) (Lepidoptera:

Choreutidae) in England, with notes on some of its parasites (Hymenop-

tera). Entomologist's Gaz. 32: 203-204.

Shaw, M. R. 198lb. Delayed inhibition of host development by the non-

paralyzing venoms of parasitic wasps. J. Invert. Pathol. 37: 215-221.

Shaw, M. R. In press. Foodplants and parasites (Hymenoptera: Ichneumonoi-

dea) of Choreutis pariana (Clerck) (Lepidoptera: Choreutidae) in

Edinburgh. Entomologist's Gaz.

328 Contrib. Amer. Ent. Inst., vol. 20, 1983

Shaw, M. R. and Askew, R. R. 1976. Ichneumonoidea (Hymenoptera) para-

sitic upon leaf -mining insects of the orders Lepidoptera, Hymenoptera

and Coleoptera. Ecol. Ent. |: 127-133.

Shenefelt, R. D. 1969. Notes on some Rogadine genera (Hymenoptera:

Braconidae). Proc. Ent. Soc. Washington 7]: 428-444.

Shenefelt, R. D. 1975. Hymenopterorum Catalogus 12: Braconidae 8, Exo-

thecinae and Rogadinae. Dr. W. Junk.

Smith, O. J., Diboll, A. G. and Rosenberger, J. H. 1955. Laboratory studies

of Pelecystoma harrisinae (Ashmead), an adventive Braconid parasite

of the Western Grape Skeletonizer. Ann. Ent. Soc. Amer. 38: 232-237.

Watanabe, C. 1962. Occurrence of Pelecystoma tricolor Wesmael in Japan

(Hymenoptera: Braconidae). Insecta Matsumurana 25: 41-43.

NOTES ON THE VARIABILITY OF MICROCTONUS AETHIOPOIDES LOAN

(HYMENOPTERA : BRACONIDAE : EUPHORINAE)

Jean-Paul Aeschlimann

335 avenue Paul Parguel, 34100 Montpellier, France

Abstract

Laboratory experiments and field observations reveal that there is a num-

ber of geographical as well as host-associated biotypes of M. aethiopoides.

Parasitoids obtained in Mediterranean France from H. postica have three, from

S. discoideus five yearly generations as compared with two only in continent-

al areas. The biotypes differ not only in their bionomics (number of genera-

tions, period of emergence, duration of pupal stage), but also in their morphol-

ogy (average number of flagellar segments) and coloration. The effect of

temperature on pupal development and coloration of female M. aethiopoides

is described and the practical implication of these observations in terms of

biological control is discussed.

INTRODUCTION

Most species of the genus Microctonus Wesmael oviposit in adult Coleop-

tera in which the larva develops as an endoparasitoid, usually sterlizing and

then killing its host. Because of their potential importance for biological con-

trol, several representatives of the genus have been recommended for use

against a number of chrysomelid and curculionid pest species (Smith 1952,

1953; Loan 1960, 1967, 1975; Drea et al . 1972; Dysart and Day 1976;

Aeschlimann 1978; Harcourt et al. 1979).

Investigations began in 1973 at the Montpellier station of CSIRO to find

efficient natural enemies of Sitona discoideus Gyllenhal (Col., Curculionidae),

a weevil that had recently become accidentally established in South-East

Australia and assumed pest proportions in fields of Medicago spp. The

euphorine Microctonus aethiopoides Loan was found to be the predominant

parasitoid of Sitona weevil in the Mediterranean region and was considered

to be the most promising biological control agent for introduction into

Australia. Rearing of the Braconid was initiated in 1974 at Montpellier and

in 1976-77 at Canberra and Adelaide after importation into Australia (Cullen

and Hopkins 1982). This paper describes some characteristics of Mediter-

ranean M. aethiopoides biotypes and compares these with findings by other

researchers in Central Europe and North America.

MATERIAL AND METHODS

The techniques used to collect and record the host weevils and their

natural enemies have been ‘described by Aeschlimann (1978, 1979).

At Montpellier, M. aethiopoides was mass-reared in polystyrene con-

tainers used for culturing Sitona (Aeschlimann 1978). The procedure was

simplified by replacing potted lucerne (Medicago sativa Linnaeus) plants with

several cut stems of lucerne (30~40 cm long) in each rearing unit. Cotton

wool was tied around the stems 6-8 cm above the cut ends. The stems were

then placed into a polystyrene tube (8 cm long, 3 cm diameter) filled with

tap water. The open end of the tube was sealed with "Parafilm" just above

329

330 Contrib. Amer. Ent. Inst.; vol. 20, 1983

the ring of cotton wool. Water was added as required through a hole in the

upper side of the tube. From June to August, the majority of cut stems de-

veloped a dense root system in the tube, making replacement of the host plant

unnecessary for several months.

The rearing units used at Canberra have been described by Cullen and

Hopkins (1982).

Adult M. aethiopoides for use in coloration studies were obtained at

Canberra by maintaining three goups of freshly spun cocoons at constant tem=

peratures of 10, 19 and 28°C and checking each group daily for emergence (the

photoregime and relative humidity in the three growth chambers were similar).

SPECIFICITY AND VOLTINISM

There are several erroneous records in the lists of hosts published by

Smith (1953), Shenefelt (1969) and Neal (1970) for M. aethiops auct. Ina

taxonomic revision of this group of euphorines, Loan (1975) showed that a num-

ber of Microctonus species had been misidentified as M. aethiops auc., and

that the only hosts of M. aethiopoides were in fact species of the genera

Hypera Schonherr and Sitona Germar (Col., Curculionidae). The following ob-

servations suggest that specificity in the field may be even greater.

In Mediterranean France, the dominant curculionid species in lucerne

growing areas are S.discoideus and the alfalfa weevil Hypera postica Gyl-

lenhal, an almost cosmopolitan pest of cultivated lucerne and also a potential

threat to Australian agriculture. The two weevils are parasitized by M.

aethiopoides, but two distinct biotypes of the parasitoid may be recognized

at sites where both species are abundant. These biotypes differ in several as-

pects of their biology and morphology, although they are not strictly specific

in terms of hosts selection. When provided with a choice under laboratory

conditions, females show a preference for one host species, but they will ovi-

posit in both hosts in a no-choice situation. However, the differing life cycles

of the weevil species contribute towards the segregation of the two different

biotypes of M. aethiopoides.

Microctonus spp. are generally passing through one generation per year,

although several Palearctic as well as Nearctic representatives of the genus

may have two, sometimes three, and exceptionally even four generations per

annum (Delucchi 1952; Smith 1952 pro errore; Jourdheuil and Chansigaud 1961;

Coles and Puttler 1963; Loan 1960, 1967, 1975; Neal 1970; Drea et al. 1972;

Dysart and Day 1976; Luff 1976; Harcourt et al. 1979; Wylie 1980). The period

over which sexually immature hosts are present in the field largely determines

the number of parasitoid generations, as such hosts inhibit the development

of first-instar Microctonus larvae. In temperate areas the curculionid hosts

of M. aethiopoides emerge during summer, and hibernate before laying most

of their eggs in the following spring. Under Mediterranean climatic conditions,

adults of S.discoideus and H. postica emerge towards the end of spring,

and aestivate before becoming sexually reproductive in early autumn (cf. Fig.

1). The bionomics of the host permits the biotype infesting S. discoideus

to complete three generations between March and June (Fig. 1), with a further

two generations from Septembr to December in parts of the French Aude, Hér-

ault and Gard départements which have a Mediterranean climate. In the same

Aeschliman: Microtonus (Braconidae) 331

localities, the biotype attacking H. postica also produces three generations

during spring, but with different emergence periods of adults as Figure |

shows; so far no M. aethiopoides adult was ever obtained from H. postica

in autumn rearings.

According to Loan (1975), a deviation of + 1 from the average number

of flagellar segments of the female antennae occurs in most species of the

genus Microctonus, Adults of M. aethiopoides obtained from the two host

species differ significantly in this respect (Table 1), with an overlap occurring

in no more than 3 % of the females.

Table 1. VARIATION IN THE NUMBER OF FLAGELLAR SEGMENTS

OF FRENCH M. aethiopoides.

Proportion of M. aethtopotdes (%) with following number of segments

| [1s | 20] a | 22 | 29 | 2 | 25 | 26 | 27 [20 | as

16.1 |. 84.51 2.4

3.6) 57i1 9 SO l7 t 336

5. OZ | Saas. fi 2esa Pibe7

OP Tau es gt PUN ile A 2 aa OM a he ©

(a) % |

Sex of

M. aethto-

Host Species

potdes

S. dtseotdeus

H. pestred

S. dtscotdeus

HH. posttea

APR. MAY ° JUN. JUL. AUG. SEP. OCT.

Fig. 1. Emergence periods of adult M. aethiopoides in cultivated lucerne

plots of Mediterranean France. (a) from S. discoideus. (b) from H. postica.

(*) beginning of new weevil generation emergence. (¥) end of overwintered

weevil generation.

PUPAL DEVELOPMENT AND COLORATION

Two biotypes of M. aethiopoides, one from a warm and dry area in

southern Morocco and the other from a cooler locality in central Greece have

been released against S. discoideus in Australia (Cullen and Hopkins 1982).

332 Contrib. Amer. Ent. Inst., vol. 20, 1983

Females from both areas show distinct seasonal variations in coloration. The

head and thorax of individuals emerging in late spring and early autumn are

orange-colored while females produced in early spring and late autumn are

blackish. Males remain black-colored throughout the season. The most

characteristic changes in coloration of female M. aethiopoides occur on the

frons and thorax (Table 2). Moroccan females tend to be somewhat lighter

in color than those of Greek origin. The biotypes differ also in the duration

of the pupal stage (Table 2). These results are supported by preliminary

electrophoretic analyses (W. J. Davidson, pers. comm.) which indicate that there

is a consistent difference in the pattern of non-specific esterases between

Moroccan and Greek specimens - despite individual varations due to age, sex,

and the temperature at which parasitoids are reared.

Table 2. EFFECT OF TEMPERATURE DURING PUPAL STAGE ON

COLORATION OF FEMALE M. aethiopoides.

Mean pupal time Coloration of female M. aethtopoides

and extremes (d)

PAU dark, lower

(26-28) 0.6 lighter

Biotype Tempera-

ture (°C)

Scutellum

black area a-

round ocellae

(10)

black

betw.

spot reddish-brown,

dark line along

upper carina

dark, median

ocellae 0.3 red-brown

(5-7)

dark line

betw. ocellae

light dark, median

0.6 reddish-

brown

dark, brown

spot on me-

dian 0.3

Moroccan black spot dark, lower

0.6 lighter

dark, upper

0.2 reddish

betw. ocellae

dark line light light light, dark

lateral spot

betw. ocellae

completely

light

light light light light, apical

0.2 lighter

DISCUSSION

Apart from Delucchi's (1952) brief account of M. emmae Delucchi in _

southern Italy, M. aethiopoides is the first species of the genus to be studied

in its Mediterranean distribution range. Mediterranean biotypes of M.

aethiopoides, passing through five annual generations, are more likely to

achieve some degree of control of S. discoideus pest populations than those

from continental areas of Central Europe and North America which only have

Aeschliman: Microtonus (Braconidae) 333

two yearly generations. Differences between biotypes originating from various

parts of the Mediterranean region also have to be considered; the shorter de-

velopmental duration of Moroccan pupae (cf. Table 2) as compared with those

of Greek origin for instance may confer a distinct advantage on the former

biotype in the field. These observations on the biotypes of M. aethiopoides

support the hypothesis (Townes 1971) that exploration for efficient biological |

control organisms should be concentrated on areas with a climate as dry and

warm, or preferably drier and warmer, than that in the area of release.

In our laboratory rearings, mortality of M. aethiopoides larvae of the

various biotypes within their hosts never exceeded 2 %. This contrasts with

the failure (through encapsulation, inability to emerge, non-establishment) to

rear and introduce M. aethiopoides elsewhere (Loan 1960, Coles and Puttler

1963, Clancy 1969, Neal 1970) probably due to the selection of a biotype poorly

adapted to the climatic conditions prevailing in the release area or primarily

parasitizing a weevil species other than the target host.

As has been suggested for the Sitona species infesting Medicago spp. in

the Palearctic region (Aeschlimann 1983), the biotypes of Microctonus para-

sitizing Sitona weevils may be undergoing speciation due to the continuous se-

lection pressure of climate, geographical isolation, and restriction to a particu-

lar host species in recently colonized territories.

ACKNOWLEDGMENTS

The author would like to thank C. C. Loan (Ottawa) and A. Roudier

(Paris) for verifying the identifications of the species dealt with, W. J.

Davidson (Canberra) for permission to mention unpublished results of biochemi-

cal analyses, G. H. L. Rothschild and R. D. Hughes (Canberra) for critically

reviewing the manuscript, J. A. Cavanaugh (Canberra) and J. Vitou (Montpellier)

for invaluable technical assistance.

7 4

RESUME

Notes sur la variabilité de Microctonus aethiopoides Loan (Hymenoptera,

Braconidae : Euphorinae)

Expériences au laboratoire et observations en nature permettent de con-

clure a l'existence de différents biotypes de M. aethiopoides. Les divergences

entre ces biotypes sont essentiellement fonction de l'origine géographique et

de I'héte préférentiel. Ceux qui évoluent en France sous un régime climatique

méditerraneen ont cing générations annuelles sur S. discoideus, mais trois

seulement sur H. postica; sous un climat tempéré en revanche, les parasitoides

de ces mémes hotes n'ont que deux générations annuelles en Europe centrale

et en Amérique du Nord. Les biotypes différent non seulement par le volti=

nisme, mais également par les périodes d'émergence des diverses générations,

la durée moyenne du stade nymphal, le nombre moyen de segments du flagelle,

la coloration corporelle des femelles et la distribution des estérases obtenue

par electrophorése. L'influence de la température sur le développement nymphal

et la coloration des femelles de M. aethiopoides est décrite. Les conséquences

pratiques de ces observations pour la lutte biologique contre les Curculionides-

hotes sont discutées.

334 Contrib. Amer. Ent. Inst., vol. 20, 1983

REFERENCES

Aeschlimann, J. P. 1978. Heavy infestations of Sitona humeralis Stephens

(Col., Curc.) on lucerne in southern Morocco. Ann. Zool. Ecol. Anim.

10: 221-225.

Aeschlimann, J. P. 1979. Sampling methods and construction of life tables for

Sitona humeralis populations (Col., Curculionidae) in Mediterranean

climatic areas. J. Appl. Ecol. 16: 405-415.

Aeschlimann, J. P. 1983. Aspects of the bionomics of the Sitona humeralis

"sroup of species", with especial consideration of S. discoideus and

S. humeralis. Ent. Exp. et Appl. (to be published).

Clancy, D. W. 1969. Biological control of the Egyptian alfalfa weevil in

California and Arizona. J. Econ. Ent. 62: 209-214.

Coles, L. W. and Puttler, B. 1963. Status of the alfalfa weevil biological con-

trol program in the Eastern United States. J. Econ. Ent. 56: 609-6ll.

Cullen, J. M. and Hopkins, D. C. 1982. Rearing, release and recovery of

Microctonus aethiopoides Loan (Hymenoptera : Braconidae) imported

for the control of Sitonahumeralis Stephens (Coleoptera : Curculionidae)

in Australia. J. Aust. Ent. Soc.

Delucchi, V. 1952. Microctonus emmae n. sp. (Hymenoptera Braconidae) paras-

sita degli adulti di Longitarsus albineus Foundr. in Puglia. Redia 37:

179-186.

stelleri (Hymenoptera : Braconidae, Euphorinae), a new parasite of the

alfalfa weevil introduced into the United States. Canad. Ent. 104: 1445-

1456.

Dysart, R. J. and Day, W. H. 1976. Release and recovery of introduced para-

sites of the alfalfa weevil in Eastern North America. USDA-ARS, Prod.

Res. Rep. 167: 1-61. |

Harcourt, D. G., Ellis, C. R. and Guppy, J. C. 1979. Distribution of Microc-

tonus aethiopoides, a parasitoid of the alfalfa weevil (Coleoptera :

Curculionidae) in Ontario. Proc. Ent. Soc. Ont. 110: 35-39.

Jourdheuil, P. and Chansigaud, J. 1961. Les parasites imaginaux des altises

inféodées au lin cultivé (Col. Chrysomelidae). Bull. Soc. Ent. Fr. 66: 219-

225,

Loan, C. C. 1960. The biology of insect parasites of the genus Sitona

Germar (Coleoptera : Curculionidae). Ph.D. thesis, Univ. Minnesota.

Pp. 1-169.

Loan, C. C. 1967. Studies on the taxonomy and biology of the Euphorinae

(Hymenoptera : Braconidae). II. Host relations of six Microctonus

species. Ann. Ent. Soc. Amer. 60: 236-240.

Loan, C. C. 1969. A summary of species of Microctonus North of Mexico

with five new species (Hymenoptera : Braconidae : Euphorinae). Proc.

Ent. Soc. Wash. 71: 404-416.

Aeschliman: Microtonus (Braconidae) 335

Loan, C. C. 1975. A review of Haliday species of Microctonus (Hym. :

Braconidae, Euphorinae). Entomophaga 20: 31-41.

Luff, M. L. 1976. The biology of Microctonus caudatus (Thomson), a braconid

parasite of the ground beetle Harpalus rufipes (Degeer). Ecol. Ent.

I: 111-116.

Neal, J. W. 1970. A study of Microctonus aethiops (Nees), a braconid para-

site of the alfalfa weevil, Hypera postica (Gyllenhal). Ph.D. thesis, Univ.

Maryland. Pp. 1-92.

Shenefelt, R. D. 1969. Braconidae |. Hybrizoninae, Euphorinae, Cosmophorinae,

Neoneurinae, Macrocentrinae. In: Hymen. Cat. (N.E.) C. Ferriére and

J. Van der Vecht (ed.), Junk Uitgev. 's-Gravenhage. Pp. 1-176.

Smith, O. J. 1952. Biology and behaviour of Microctonus vittatae Muesebeck

. (Braconidae) with descriptions of its immature stages. Univ. Calif. Publ.

Ent. 9: 315-343.

Smith, O. J. 1953. Species, distribution, and host records of the braconid

genera Microctonus and Perilitus... Ohio 4. Scis.53: 173-178.

Townes, H. 1971. Ichneumonidae as biological control agents. Proc. Tall Timb.

Conf. Ecol. Anim. Contr. Habit. Manag. Pp. 235-248.

Wylie, H. G. 1980. Colour variability among females of Microctonus

vittatae (Hymenoptera : Braconidae). Canad. Ent. 112: 771-774.

TAXONOMIC AND BIOLOGICAL STUDIES OF DIVERSINERVUS CERVANTESI

(GIRAULT) (HYMENOPTERA : ENCYRTIDAE), A PRIMARY PARASITE

OF SOFT SCALE INSECTS

David Rosen and Amos Alon

The Hebrew University, Faculty of Agriculture

Rehovot, Israel

Abstract

Diversinervus cervantesi (Girault) (Hymenoptera: Encyrtidae) is a pri-

mary, gregarious endoparasite of soft and wax scale insects (Homoptera:

Coccidae), recorded from Australia, Malaysia and Israel. The species is

deuterotokous, with brachypterous females and rare macropterous males. The

adult female, the hitherto undescribed male and the developmental stages are

described and figured.

The eggs of D. cervantesi are laid into the hind gut of the host. De-

velopment takes 2] days at 28°C, 28 days at 24°C, 44 days at 20°C; the

threshold of development is 12.5°C and the thermal constant is 327.8 day-

degrees. The females exhibit high longevity (average 45 days at 26°C) and

high fecundity (average 187 progeny). They are very sensitive to low relative

humidity, whereas temperatures in the range of 24-32°C to not have a pro-

nounced effect on their longevity.

Two informal species groups are proposed for the 10 valid species pres-

ently included in the genus Diversinervus Silvestri: The elegans group, com-

prising desantisi Compere, elegans Silvestri, madgaoensis Hayat, Alam and

Agarwal, paradisicus (Motschulsky), scutatus Compere, silvestrii Waterston

and stramineus Compere, in which the females have tufts of bristles on both

the mesoscutum and scutellum, the head of the males is metallic, the eggs

are banded and the first-instar larvae have open tracheal systems; and the

smithi group, comprising cervantesi (Girault), masakaensis Compere and

smithi Compere, in which the females lack the mesoscutar tuft, the head of

the males is yellow, the eggs are unbanded and the first-instar larvae have

closed tracheal systems.

A revised key to the species of Diversinervus is presented.

INTRODUCTION

Diversinervus cervantesi (Girault) (Hymenoptera: Encyrtidae) is a pri-

mary gregarious endoparasite of various soft and wax scale insects (Homoptera:

Coccidae). It is an unusual species, with brachypterous females and rare

macropterous males which, in spite of its limited mobility, appears to be

rather widespread, having been recorded so far from Australia, Malaysia and

Israel. Inasmuch as it was discovered in Israel for the first time only a few

years ago, we assume that it may have invaded the country fairly recently,

presumably with host material. The taxonomic and biological studies reported

herein were conducted as part of a long-term research project on the hymen-

opterous parasite fauna of Israel.

The genus Diversinervus Silvestri is closely related to Cheiloneurus

Westwood, a genus of injurious hyperparasites, and was therefore at first also

336

Rosen & Alon: Diversinervus (Encyrtidae) 337

thought to be hyperparasitic. However, Compere (1931) has shown that the

type-species, D. elegans Silvestri, in fact develops as a primary parasite of

coccids. All known members of the genus are presently believed to be pri=-

mary parasites, and some of them have been utilized in the biological control

of coccid pests (Bartlett 1978).

Diversinervus was initially separated from Cheiloneurus by having con-

spicuous tufts of bristles on both the mesoscutum and scutellum of the female.

This, however, proved to be an unreliable generic character. Several bona fide

members of Diversinervus, including cervantesi, resemble Cheiloneurus in

lacking the mesoscutas tuft (Fig. 1). As pointed out by Compere (1940), the

main diagnostic character by which Diversinervus may be distinguished from

Cheiloneurus and other related genera is the peculiar structure of the female

head, with the frontovertex expanded into a frontal ledge and the face meet-

ing it in a sharp angle (Fig. 1). Also, in most species of Diversinervus, the

marginal vein of the female fore wing is relatively short, subequal in length

to the stigmal vein, whereas in Cheiloneurus it is considerably longer.

Fig. 1. Diversinervus smithi Compere, Female head and thorax

(From Compere, 1940)

Diversinervus cervantesi (Girault), n. comb.

Cheiloneurus cervantesi Girault, 1933. Some Beauties Inhabitant Not

of Commercial Boudoirs but of Nature's Bosom, Notably New Insects. Priv.

Publ., Brisbane, p. 4.

Girault (1933) described this species as follows:

"Cheiloneurus cervantesi. Subapterous, wing | fuscous at apex. Like

Diversinervus genotype but ocelli absent(?), hind margin pronotum with thick

black bristles like those of hair-tuft, thicker mesad; metatarsus white. Discal

cilia absent, no stigmal or postmarginal vein. Ex Pulvinaria. Nambour, Feb.

A. R. Brimblecombe."

REDISCRIPTION

Female (Figs. 2-20): Head in lateral view (Fig. 2), triangular; in dorsal

view (Fig. 3), somewhat wider than the thorax (1.0 -1.1), somewhat wider than

long (1.10 -1.25); frontovertex deeply emarginate posteriorly, mostly parallel-

338 Contrib. Amer. Ent. Inst., vol. 20, 1983

sided, 3.5 -3.9 times as long as wide, 0.22 ~0.26 width of head, smooth, bear-

ing 2 longitudinal rows of fine setae, with 2 pairs of longer, coarser, black

setae along occipital margin, on either side of indentation. Ocelli small, in

an equilateral triangle, the posterior pair about twice their own diameter from

orbits, 5 diameters from occipital margin. Eyes large, 1.4 -1.6 times as long

as wide, bordered by a row of fine setae and bearing short, sparse inter-

ommatidial setae. Face and cheeks reticulate; toruli (Fig. ll) subtriangular,

their bases at about one-third the distance between clypeal and frontal mar-

gins; scrobes meeting anteriorly. Mouthparts (Figs. 4-6): Mandibles triden=

tate, 2.0-2.5 times as long as wide, bearing about 7 setae, the denticles sub-

equal. Maxillary palpi 4-segmented, the distal segment about twice as long

as the penultimate (1.85 -2.20), bearing several subapical setae, the longest

somewhat longer than the segment itself. Labial palpi 3-segmented, second

segment the shortest and wider than long, the distal one bearing several setae.

Antennae (Figs. 7-ll): Radicle stout, longer than wide, bearing several

finger-like sensilla. Scape moderately expanded antero~ventrally, about 2.7

times as long as wide, 2.4-2.8 times longer than the pedicel, 1.10-1.25 times

longer than the club. Pedicel conical, about twice as long as wide (1.8 - 2.3),

about twice longer and one-fifth to one-third wider than the first segment

of the funicle. Funicular segments I-III as long wide or somewhat longer,

segments IV-VI as long as wide; segments I-IV subequal in length, V about

one-third longer (1.23 -1.43) and one-fourth wider (1.20 -1.35) than IV, VI some-

what larger than V; setae on segments I-III about as long as the segments.

Club about 3 times as long as wide (2.6 -3.3), equal in length to the preced-

ing 4 segments and one-third to one-half of segment II combined, about one-

fifth wider than segment VI (1.08 -1.32); all 3 club segments bearing

longitudinal sensilla, funicle V and VI bearing one sensillum.

Thorax (Figs. 2, 12-13): As in many other brachypterous encyrtids, ap-

ears rather undeveloped, widening posteriorly, with small mesonotal and

metanotal sclerites and relatively large pronotum, tegulae and mesopleura.

Pronotum large, somewhat wider than long, widening posteriorly, overlapping

the anterior margin of mesoscutum, reticulate, bearing 3 transverse rows of

about 20 setae each, those of the posterior row black, longer and coarser sub-

medially. Mesoscutum convex, as long as the pronotum and somewhat longer

than wide, mostly reticulate-strigose, lacking a tuft of bristles but with a

smooth area latero-posteriorly bearing one lateral row and 3 irregular posterior

rows of silvery setae. Axillae meeting mesally, reticulate, each bearing 1-2

pale setae. Scutellum convex, rather narrow, as long as the mesonotum and

one-fourth to one-third longer than wide, occupying only one-third to two-

fifths width of thorax (0.34 -0.40), reticulate, bearing a dense tuft composed

of 2 submedian longitudinal rows of 4-6 black, coarse bristles each,

surrounded by several shorter setae, and a pair of longer erect subapical

bristles. Tegulae large, triangular, reticulate, each bearing 4 setae. Meso-

pleura large, convex, smooth, readily visible in dorsal view. Metanotum

transverse, somewhat arcuate, 4-5 times as wide as long, with coarse oblique

striation. Propodeum (Fig. 12) almost as wide as the thorax, about 3 times

as wide as long, mostly smooth, with deep transverse furrows along anterior

and posterior margins and some coarse striation antero-laterad of spiracles,

the latter areas bearing a single fine seta.

Rosen & Alon: Diversinervus (Encyrtidae) 339

Gaster (Figs. 2, 14-15) about one and one-half times as long as wide,

somewhat shorter than the thorax and about as wide; first tergite ring-like,

the second (= abdominal tergite III) large, reticulate, bearing what appear to

be numerous discoid sensilla; subsequent tergites reticulate, bearing sparse

setae; epipygium large, smooth, bearing sparse setae. Cerci situated at

basal one-third of gaster, bearing 3 long setae and 2 short setae. Ovipositor

about 0.8-0.9 length of gaster, slightly exserted, the hypopygium covering only

its basal one-third or so.

Legs (Figs. 2, 16, 17) well developed. Mid-tibial spur somewhat (up to

1.14 times) longer than the corresponding basitarsus.

Wings (Figs. 12-13, 18-20) greatly reduced, variable. Fore wing about

as long as width of thorax, usually not reaching beyond second gastral tergite,

narrow, about 6.5 times as long as wide (6.3 -6.8); submarginal vein bearing

3-4 fine setae proximally, somewhat expanded distally and bearing 6-8 setae,

usually reaching margin at apex of wing, with an apical bristle about as long

as width of wing; a few fine discal setae along anterior margin and at postero-

distal corner, marginal fringe usually absent; sometimes the wing is somewhat

expanded beyond venation (Fig. 19), the distal area bearing discal setae and

a short marginal fringe. Hind wing usually very narrow, considerably shorter

than the fore wing, the vein reaching apex of wing, with 3 apical hamuli; oc-

casionally (Fig. 20), the wing is larger, expanded beyond venation, bearing

numerous discal and some marginal setae.

General coloration orange-yellow. Eyes dark purple, ocelli reddish brown.

An arcuate dark brown stripe on face and cheeks, at level of antennal toruli.

Distal one-third of mandible dark brown. Antenna concolorous with body,

ventral margin of scape and dorsal margins of pedicel and funicle segments

I~V lined with dark brown; first club segment suffused with brownish distally,

second and third segments dark brown. Mesoscutum with a light metallic

cross-band on posterior one-third, bearing silvery setae; scutellum somewhat

paler yellow, with a pair of brown blotches posteriorly, on either side of

tuft; propleura, prosternum and mesosternum lined with dark brown. Propodeum

with brown blotches at postero-lateral corners. Gaster lined with dark brown

on margins, with a semi-circular transverse dark brown blotch centrally, be-

tween cerci; cercal plates dark brown. Legs concolorous with body, the follow-

ing parts translucent white: fore and hind coxae, all trochanters, basal two-

thirds of fore femur, three-fourths of middle femur and one-third of hind

femur, base of middle and hind tibia, and most of hind tarsus; dark brown on

‘middle coxa, a spot near base of middle and hind tibia, and all pretarsi.

Fore wing hyaline, marked with dark brown on proximal part of vein, on frenal

fold at proximal part of posterior margin, and at apex, including distal part

of vein; when somewhat expanded beyond venation, that apical part is hyaline,

the dark brown area then forming an oblique cross-band from apex of vein

to posterior margin. Hind wing hyaline.

Length: 1.20-1.56 mm.

Male (Figs. 21-27): Differing markedly from the female in the structure

of head, antennae, thorax and wings, as well as in general coloration.

Head in lateral view, lenticular; in dorsal view, nearly as wide as the

thorax (0.83 -0.95), almost 3 times as wide as long (2.6 -2.8), delicately

340 Contrib: “Amer. Ent. insta; vol. 205 1983

reticulate; frontovertex broad, about one-third wider than long, about half

width of head (0.47-51). Ocelli large, in an obtuse triangle (100°), the

posterior pair about their own diameter from orbits, half a diameter from

occipital margin. Eyes smaller than in the female. Toruli situated at about

middle of face; scrobes meeting anteriorly. Mouthparts as in the female.

Antennae (Fig. 21) filiform. Radicle stout, 1.3 -1.6 times as long as

wide. Scape about 3 times as long as wide, distinctly shorter than the club.

Pedicel about one and one-half times as long as wide (1.40 -1.56),-consider-

ably shorter than the first segment of the funicle (0.5-0.7). All funicular

segments elongate, subequal in width, bearing whorls of long setae, segments

V-VI bearing several longitudinal sensilla; segments II-V subequal in length,

about 3 times as long as wide, I and VI somewhat shorter, about 2.5 times

as long as wide. Club undivided, elongate, 4-5 times as long as wide, about

three-fourths longer and somewhat wider than the preceding segment, bearing

long setae and several longitudinal sensilla.

Thorax (Fig. 22) well developed, convex, rather uniformly reticulate dor-

sally, the scutellum reticulate-strigose mesally. Pronotum conical, bearing

2-3 transverse rows of setae. Mesoscutum rather broad, somewhat wider than

long, bearing numerous sparse setae. Axillae separate, their apices not meet-

ing mesally, each bearing 1-2 setae. Scutellum triangular, somewhat shorter

than the mesoscutum and about as long as wide, occupying one-half to two-

thirds width of thorax (0.56 -0.65), bearing 2 sparse, longitudinal rows of

6-7 setae each, the posterior pair longer than the rest. Tegulae triangular,

smaller than in the female, each bearing several setae. Mesopleura large,

smooth, not visible in dorsal view. Metanotum V-shaped, about 5-7 times as

wide as long, with coarse oblique striation. Propodeum (Fig. 22) about as

wide as the thorax, 3-4 times as wide as long, mostly smooth, with some

transverse striation along anterior and posterior margins and antero-laterad

of spiracles.

Gaster (Fig. 23) trapezoidal, about one and one-half times as long as

wide, about three-fourths length of thorax (0.66 -0.81) and about as wide,

largely smooth, bearing sparse setae. Cerci as in the female. Genitalia

(Figs. 23, 24) 3.4-5.5 times as long as wide, about two-fifths length of

gaster (0.35 -0.46), three-fifths to three-fourths length of middle tibia (0.58 -

0.76), exserted to about one-third length; digital sclerites about one-fifth the

combined length of aedeagus and slender apodemes (0.17 -0.23), bearing a

single strong claw.

Legs (Fig. 25) similar to those of female. Mid-tibial spur about half

as long as the corresponding basitarsus.

Wings (Figs. 26, 27) well developed. Fore wing about as long as the

entire body, 2.5 times as long as wide (2.3-2.9), rather uniformly setose

distad ot speculum, the setae proximad of speculum somewhat longer and

sparser; marginal fringe not exceeding one-tenth width of disc; submarginal

veing somewhat widened on distal one-third, bearing 10-12 setae and numerous

bullae; marginal vein twice as long as wide, subequal in length to the stigmal

and postmarginal veins, bearing several setae; stigmal vein about one and one-

half times as long as wide, widening distally, bearing 3 discal sensilla in a row

at apex; postmarginal vein tapering toward apex, bearing several setae. Hind

Rosen & Alon: Diversinervus (Encyrtidae) 341

wing about 4-5 times as long as wide, two-thirds length of fore wing, uniform-

ly setose except at base; marginal fringe not exceeding one-third width of

disc; venation reaching to about two-thirds length of wing, the marginal vein

bearing 3 apical hamulli.

Coloration considerably darker than in the female. Head orange-yellow,

with a pair of small dark brown blotches on occiput, on either side of foramen.

Eyes and ocelli purplish black. Antennae uniformly brown, scape somewhat

paler, pedicel marked with blackish dorso-basally, flagellar setae dark brown.

Pronotum dark metallic centrally, orange-yellow laterally; mesonotal sclerites

dark metallic, with green reflections, sides of mesoscutum orange-yellow,

lateral and posterior margins of axillae narrowly lined with orange-yellow;

tegulae brownish, mesopleura brown; metanotum brown; venter of thorax

orange-yellow. Propodeum mostly brown, the antero-lateral corners orange-

yellow. Gaster entirely black. Legs mostly pale yellow; hind tibia faintly-

marked with brownish, fore tarsus brown, distal segment of middle and hind

tarsi brownish. Wings hyaline, the venation blackish; submarginal vein of fore

wing interrupted subapically by a short hyaline portion.

Length: 0.8-1.0 mm.

Redescribed from numerous male and female specimens, insectary-reared

on Coccus hesperidum L. on citron melon, Citrullus vulgaris var. cit

oides, Rehovot. The stock had originated from 3 females, reared ex

Pulvinaria mesembryanthemi (Vallot) on Mesembryanthemum edule, Rehovot,

Israel.

Notes: The type series of cervantesi was received on loan from the

Queensland Museum, Brisbane, Australia. It consists of 2 female syntypes,

mounted in balsam on one slide, labeled "Cheiloneurus cervantesi Gir. F Types"

in A. A. Girault's handwriting. The body of one female lies under one cover

glass, a head and the dissected parts of another female under another cover

glass. We assume that the undissected head belongs to the undissected body,

but this is by no means certain. Designation of a lectotype is therefore un-

desirable.

The Israel material described herein appears to be conspecific with the

syntypes. Some of the syntype antennae seem to be somewhat more exten-

sively pigmented, with the first club segment mostly - or entirely — fuscous,

but antennal pigmentation appears to be variable in the type series (Figs. 7,

8) and cannot be used as a diagnostic character. Specimens received for

identification from Malaysia (recorded as Diversinervus sp. by Chua, 1976)

were also found to be conspecific.

Although Girault (1933) compared cervantesi to the generotype of

Diversinervus, he referred this peculiar species to the genus Cheiloneurus

but failed to state his reasons for doing so. His decision was probably based

on the absence of a mesoscutar tuft. This species is a bona fide member of

the genus Diversinervus. It appears to be closely related to D. smithi

Compere, a macropterous species described by Compere (1940) from South

Africa. It resembles smithi in most aspects of structure, chaetotaxis, sculp-

ture and coloration, differing mainly in the reduced wings and thoracic ter-

gites of the female. Although Girault's tentative reference to the absence

of ocelli in cervantesi was erroneous, the ocelli of the female are indeed

342 Contrib. Amer: Ent. Inst., vol. 20; 1983

Figs. 2-6. Diversinervus cervantesi (Girault), female: 2, Lateral view

of entire specimen (SEM). 3, Head, dorsal view (SEM). 4-6. Mouthparts

(4 = syntype; 5 = SEM).

Rosen & Alon: Diversinervus (Encyrtidae) 343

Figs. 7-13. Diversinervus cervantesi (Girault), female: 7-10, Antennae

(7, 8 = syntype; 10 SEM). 11, Base of antenna (SEM). 12, Thorax, propo-

deum and fore wings (SEM). 13, Scutellum, axillae and fore wings (SEM).

Contrib. Amer. Ent.-Inst., vol. 20, 1983

344

(SEM).

female: 14,15, Gaster

tesi(Girault),

17, Mid tibial spur and basitarsus

20, Hind wing, unusually developed.

iversinervus cCefvan

16, Middle leg.

Figs. 14-20. D

SEM).

eee

(14

18-19, Fore wings.

Rosen & Alon: Diversinervus (Encyrtidae) 345

Figs. 21-27. Diversinervus cervantesi(Girault), male: 21, Antennae. 22,

Thorax and propodeum (SEM). 23, Gaster (SEM). 24, Genitalia. 25, Strigil

of fore leg (SEM). 16, Fore wing. 27, Hind wing.

346 Contrib. Amer. Ent. :inet., vol. 20, 1983

smaller and less conspicuous than those of smithi, and the posterior pair is

situated somewhat farther from the occipital margin than in that species.

Also, funicular segment V islined with blackish in cervantesi, entirely yellow

in smithi; the middle coxa is brown in cervantesi, white in smithi; and the

pronotal setae appear finer, less conspicuous, in the female of smithi. The

two species are almost indistinguishable in the male sex. The males of

smithi appear somewhat more extensively pigmented than those of

cervantesi, the mesoscutum with less clearly defined yellow areas laterally

and with strong violaceous reflections on the posterior half. The frontovertex

appears somewhat narrower in the male of smithi.

As shown below, D. cervantesi also differs from D. smithi in the site

of oviposition in the host, and in the morphology of the first-instar larva.

The two species also appear to differ in their mode of reproduction: D.

cervantesi is uniparental, whereas D. smithi is presumably biparental

(Flanders 1939).

DEVELOPMENTAL STAGES

Egg (Figs. 28-31): The egg of D. cervantesi is unbanded (Maple 1947),

similar to that of D. smithi (Flanders 1952). The ovarian egg (Fig. 28) is

elongate, the dorsal aspect concave, with a narrow bulb, shorter than the egg

proper and separated from it by a short, indistinct stalk.

Unlike D. smithi, which reportedly oviposits in the suboesophageal

ganglion of its host (Flanders 1952), D. cervantesi lays its eggs in the host's

hind gut (Fig. 29). The deposited egg is banana-shaped, with the short, col-

lapsed remnants of the bulb and stalk at its anterior end (Figs. 30,°31).. It

lies freely in the gut and does not change much in size after deposition. At

28°C, the mandibles and midgut of the developing embryo become evident on

the fourth day after deposition (Fig. 31).

Larva (Figs. 32-40): The first-instar larva (Figs. 32, 33) leaves the hind

gut soon after hatching. Unattached to the egg chorion, it floats freely in

the body fluids of the host. It is orange-yellow, distinctly caudate, and lacks

the cephalic vesicles reported in D. smithi (Flanders 1952). Like other

encyrtids with unbanded eggs, the first-instar larva of D. cervantesi has a

closed tracheal system (Maple 1947). The average size of this instar increases

during its development, from 333 x 57 to 600 x 105 #. The mandibles (Fig.

33) are sickle-shaped, 8.5 # long, with a bifid base.

The fully-developed larva (Figs. 34-40) is brown-yellow, hymenopteriform,

about 1.5 mm long. It has an open, peripneustic tracheal system, with 9

pairs of functional spiracles present in thoracic segments II, III and abdominal

segments I-VII (Figs. 35, 36). Cephalic skeleton as in Figure 37 (terminology

adapted from Vance and Smith, 1933). The mandibles (Figs. 37-39) are 45 4

long, with a strong, straight tooth and a subquadrate base.

Prior to pupation, the larva expels several ovoid, dark brown meconia

which line the walls of its cell inside the host, and turns translucent, whitish

in color. This short prepupal stage is followed by a molt into the pupal stage.

Pupa (Figs. 41-44): The newly-formed pupa is pale, translucent. It lies

dorsal-side-up in its cell, surrounded by the meconia. In the female pupa,

Rosen & Alon: Diversinervus (Encyrtidae) 347

Figs. 28-34. Diversinervus cervantesi (Girault), developmental stages:

28, Ovarian egg. 29, Two eggs in host's gut. 30, Egg, 2 days old. 31, Egg,

4 days old. 32, First instar larva. 33, Mandibles of first instar larva.

34, Fully developed larva (SEM).

348 Contrib. Amer. Ent. Inst., vol. 20, 1963

Figs. 35-40. Diversinervus cervantesi (Girault), developmental stages:

35,36, Respiratory system of fully developed larva. 37, Cephalic skeleton of

fully developed larva (ep = epistoma; hyp = hypostoma; ipr = inferior pleuro-

stomal ramus; m = mandible; ma = mandibular acron; oes = oesophagus; pc =

preoral cavity; pl = pleurostoma; sd = salivary dut; spr = superior pleurostomal

ramus; ten = tentorium; terminology adapted from Vance and Smith, 1933).

38, Mandibles of fully developed larva. 39, Cephalic region of fully developed

larva, showing mandibles (SEM). 40, Two larvae in host.

Rosen & Alon: Diversinervus (Encyrtidae) 349

Figs. 41-44. Diversinervus cervantesi : Developmental stages:

41, Female pupa, ventral view (SEM). 42, Female pupa, dorsal view (SEM).

43, Three female pupae in host. 44, Two male pupae in host.

350 Contribe-Amer. Ents Inst vol. 20, 1983

pigmentation is first evident in a triangular area on the gaster, followed by

the eyes, the mandibles and the rest of the body. In the male pupa, dark

cross-bands first appear on the gaster, widening gradually and merging on the

posterior segments, then anteriorly; the eyes become pigmented at about the

same time; darkening of the thorax commences somewhat later, at the an-

terior end.

BIOLOGY

D. cervantesi is a primary, gregarious endoparasite of soft and wax

scale insects (Homoptera: Coccidae). It was recorded in Australia from

Pulvinaria sp. (Girault 1933), in Malaysia from the hemispherical scale,

Saissetia coffeae (Walker), on coffee (Chua 1976). In Israel, it has been

reared from the mesembryanthemum scale, Pulvinaria mesembryanthemi

(Vallot), on Mesembryanthemum edule, from the hemispherical scale on coffee,

from the Mediterranean black scale, Saissetia oleae (Olivier), on citrus and

oleander, and from the Florida wax scale, Ceroplastes floridensis Comstock,

on English ivy (Ben-Dov 1970, Kfir and Rosen 1980a, and in the present study).

In the laboratory, we reared it successfully on the brown soft scale, Coccus

hesperidum L., and that species served as host in all our biological studies.

Duration of Development at 28°C

Female wasps were allowed to oviposit for 24 hours in brown soft scales

on a citron melon. The parasitized hosts were then kept in an incubator at

28°C, and a sample was dissected daily to determine the stage of development

of the developing parasites.

7, l=

0 5 10 15 20 29

DAYS

Fig. 45. Diversinervus cervantesi: Development of immature stages at 28°C.

The results are presented in Figure 45. Egg incubation was relatively

long, about one-fourth of total development, the eggs hatching 5-6 days after

deposition. Larval development took about one week, pupation occurring

Rosen & Alon: Diversinervus (Encyrtidae) 3511

12-15 days after oviposition. The pupal period was rather short, occupying

only about one-third of the total; complete imaginal pigmentation was evident

as early as on the 18th day, and emergence occurred 19-25 days after oviposi-

tion.

Effect of Temperature on Rate of Development

Female wasps were allowed to oviposit for 24 hours, and the citron

melons bearing the parasitized hosts were then placed in incubators at constant

temperatures of 20, 24, 28 and 32°C. Emergence of parasite progeny was

recorded daily.

Table I. Duration of development of D. cervantesi F at

constant temperatures.

Duration of development (days)

Temperature

(°C) ‘ Range Average + SD

20 664 39-50 43.84 + 2.24

24 352 26-33 28.09 + 1.49

28 1190 19-27 21.08 & 1.30

32 0 _ ea

As expected, the duration of development was progressively shorter at

higher temperatures (Table I). No emergence occurred at 32°C. However,

when hosts containing pupae of the parasite were exposed to 32°C, many of

the parasites completed their development and emerged, indicating that this

temperature is probably lethal mainly to the early developmental stages of

D. cervantesi.

The equilateral hyperbola equation, based on the assumption that the

product of developmental time and the corresponding temperature above a cer-

tain threshold is a constant (the thermal constant) for any given species, is

a convenient way of expressing the effects of temperature on the duration of

development of invertebrates (Bodenheimer 1926, 1958). The hyperbola equation

for D. cervantesi was calculated as follows: Rates of development at 20,

24 and 28°C were obtained as the reciprocals of the actual developmental

periods recorded at these temperatures; the linear regression (y = a+bx) of

developmental rate on temperature was then calculated by the method of least

squares; the parameters of the equilateral hyperbola (thermal constant, ;

and developmental threshold, c) were then obtained from the identities TAC =

L/D 81s afe,

The regression of development rate on temperature and the corresponding

equilateral hyperbola for D. cervantesi are presented in Figure 46. The de-

velopmental threshold was 12.47°C and the thermal constant 327.76 day-

degrees.

352 Contrib. Amer. Ent. Inst., vol. 20, 1983

Y=0.00305x-0.0380

0.04

150

0.03

100

0.02

RATE OF DEVELOPMENT (1/t)

DURATION OF DEVELOPMENT (days)

0.01 t (x-12.47) =327,76

12 16 28

20 24

TEMPERATURE (°C)

Fig. 46. Diversinervus cervantesi: Effect of temperature on rate and

duration of development.

Reproduction and Fecundity

D. cervantesi is a deuterotokous species, unmated females usually giving

rise to female progeny, occasionally to rare males. Of a total of 3278 speci-

mens reared in the present study, 59, or 1.8 percent, were males.

Rearing temperatures in the range of 20-28°C did not affect the sex

ratio even after several generations of continuous exposure. Some females ex-

posed to 32°C during pupal development did not lay any eggs, whereas others

started to do so only after 7 days at a lower temperature and they, too, pro-

duced the usual sex ratio.

Development is definitely gregarious. In the laboratory, only about 10

percent of the parasitized hosts examined yielded a single parasite, whereas

about 50 percent yielded 4 parasites or more. Up to 13 parasites have been

recorded from a single host. Several adults may emerge from one exit hole

in the host's dorsum.

Similar data were obtained from dissection of parasitized hosts. Under

normal conditions, practically all the eggs laid in a host develop to maturity.

Only under extreme crowding did the number of eggs per host greatly exceed

the number of emergent progeny, and dissection of such hosts indicated that

lop)

PROGENY PER FEMALE

200

150

PROGENY PER FEMALE

ro)

Fig. 47-48.

6 females). 48,

Rosen & Alon: Diversinervus (Encyrtidae) 353

nee is id ai [ ls aa it ae

10 20 30 40

DAYS ca 60

DAYS

Diversinervus cervantesi:

47, Daily fecundity (averages of

Cumulative fecundity (averages of 6 females).

354 Contrib. “Amer. Geto insts vole, 1983

supernumerary parasites are eliminated in the egg or early larval stage. Prog-

eny emergence was therefore used as the criterion for effective fedundity.

To assess the fecundity of D. cervantesi, 6 females were allowed to

oviposit individually in brown soft scales on citron melons, at 26+1°C and

under constant illumination. (Preliminary observations had indicated that in

darkness the fecundity is considerably lower). They were transferred daily

to fresh hosts during the first week, every 1-3 days thereafter. Progeny emer-

gence was recorded daily. The results are presented in Figures 47, 48.

The ovipositing females exhibited high longevity (20-67 days, average

45.5) and high fecundity (85-307 progeny per female, average 187). The

species is synovigenic: Oviposition usually commenced on the seond day after

emergence, or near the end of the first day, and continued throughout most

of the female's adult life span. On the average, 72 percent of the progeny

were produced during the first half of the female's adult life.

Effect of Relative Humidity on Adult Life Span

Freshly-emerged females of D. cervantesi were confined in clear plas-

tic vials (diameter 43 mm, height 48 mm), the inner walls of which were

smeared with a little honey. The mouth of each vial was covered with tissue

paper and sealed with Permagum ® * to the mouth of a similar vial containing

either dry P505 or a solution of 62.5 g KOH in 100 ml water, producing a rela-

tive humidity of 0 or 50 percent, respectively, in the sealed chambers thus

formed (Peterson 1964). The vials were kept at 28°C, in the dark, and mor-

tality of the wasps was recorded every few hours during the first two days,

daily thereafter.

Table Il. Longevity of adult D. cervantesi females at 0

and 50% relative humidity and 28°C.

RH 2 Longevity Days to 50%

(%) (days) mortality

0 44 1-4 Ls

50 40 2-42 17.0

The results (Table II, Fig. 49) indicate that the females of D. cer-

vantesi are very sensitive to low relative humidity (see Bartlett 1962).

*A non-toxic, odorless sealing compound distributed by Virginia Chemicals,

West Norfolk, VA.

Rosen & Alon: Diversinervus (Encyrtidae) 255

100

(ep)

SURVIVAL

*lo

DAYS

100

*lo SURVIVAL

ron)

eo)

La)

0 10 15

DAYS

Figs. 49-50. Diversinervus cervantesi: 49, Longivity of adult females at

O and 50% RH and 280C. 50, Longivity of adult females at various constant

temperatures.

356 Contrib. Amer. Ent. Inst., vol. 20, 1983

Effect of Temperature on Adult Life Span

Freshly-emerged females of D. cervantesi were confined in closed

chambers, as described above, at 50 percent relative humidity with honey as

food. The vials were kept at 24, 28 or 32°C, in the dark, and mortality of

the wasps was recorded daily.

Table Ill. Longevity of adultD. cervantesi females at various

constant temperatures.

Temperature = Longevity Days to 50%

Sek, 2 (days) mortality

24 20 2-29 1.7

28 40 2-24 Li

32 47 2-24 Lise

The results (Table Ill, Fig. 50) indicate that temperatures in the range

of 24-32°C do not have a pronounced effect on the longevity of D.

cervantesi females.

Although the wasps in this experiment were kept in the dark, which would

presumably tend to reduce their activity and thus prolong their life span, even

at 24°C their longevity in the absence of hosts was less than half that of the

ovipositing females at 26°C and under constant illumination, as described

earlier. This difference might be ascribed to predatory host-feeding which,

however, was not observed in the present study, or to feeding on the honeydew

produced by the hosts, etc. Similar observations have been reported for other

coccid-inhabiting encyrtids (e.g., Kfir and Rosen 1980b).

SYNOPSIS OF THE SPECIES

Genus DIVERSINERVUS Silvestri

Diversinervus Silvestri (1915: 301-304). [Publ. Feb. 20, 1915].

Type-species: Diversinervus elegans Silvestri.

Cheiloneuroides Girault (1915: 96, 122). [Publ. June 4, 1915].

Type-species: Cheiloneuroides bicristatus Girault.

Taxonomy: Girault (1917: 37); Compere (1931: 233-235; 1938: 322-323;

1940: 407); Trjapitzin (1961: 671); Hayat et al. (1975: 39-41); Noyes (1980: 193).

The genus presently includes 10 valid species, which are listed below.

An additional species, D. intermedius Hayat, Alam and Agarwal (1975: 43-44),

described from a single female, ex Chloropulvinaria psidii (Maskell), India,

has not been included here. From the description, it appears to be a small

specimen of D. elegans.

Rosen & Alon: Diversinervus (Encyrtidae) 357

Diversinervus cervantesi (Girault)

Cheiloneurus cervantesi Girault (1933: 4).

Distribution: Australia, Malaysia, Israel.

Hosts: Coccidae: Ceroplastesfloridensis Comstock; Coccus hesperi<=

dum L. (in lab.); Pulvinaria: mesembryanthemi (Vallot); Pulvinaria Sp.;

Saissetia coffeae (Walker); S. oleae (Olivier).

Diversinervus desantisi Compere

Diversinervus de santisi Compere (1931: 239-241).

Distribution: Eritrea.

Host: Coccidae: Pulvinaria sp.

Diversinervus elegans Silvestri

Diversinervus elegans Silvestri (1915: 304-305).

Cheiloneuroides bicristatus Girault (1915: 96-97).

Diversinervus meridionalis Compere (1931: 237-239).

Taxonomy: Girault (1917: 37); Compere (1931: 244-245; 1937: 43-44);

Annecke (1964: 15); Rosen (1966: 61); Beardsley (1976: 189, 211).

Biology: Compere (1931); van den Bosch et al. (1955); Bartlett and

Medved (1966); Rosen (1967); Bartlett (1978); Kfir and Rosen (1980a).

Distribution: Eritrea, Kenya, South Africa, Israel, Australia, Hawaii,

Brazil, Peru, California (introd.).

Hosts: Coccidae: Ceroplastesfloridensis Comstock; Ceroplastes sp.;

Coccus hesperidum L., C. pseudomagnoliarum (Kuwana) (in lab.); Gascardia

brevicauda (Hall); G. destructor (Newstead); Marsipococcus proteae (Brain);

Parasaissetia mangiferae (Green); Pulvinaria mesembryanthemi (Vallot);

Saissetia coffeae (Walker); S. oleae (Olivier); S. persimilis (Newstead);

Saissetia spp.

For additional references see Peck (1963), Annecke and Insley (1971).

Diversinervus madgaoensis Hayat, Alam & Agarwal

Diversinervus madgaoensis Hayat, Alam and Agarwal (1975: 41-42, 44).

Distribution: India, Philippines.

Hosts: Coccidae: Ceroplastes actiniformis Green; C. ceriferus

(Fabricius); Pulvinaria sp.

Note: Specimens of D. madgaoensis ex C. ceriferus, Banaras, India and

ex Pulvinaria sp., Manila, Philippines, stand in the USNM.

Diversinervus masakaensis Compere

Diversinervus masakaensis Compere (1940: 407-408).

Distribution: Uganda.

Host: Coccidae: Saissetia oleae (Olivier).

358 Contrib. Amer. Ent. Inst., vol. 20, 1983

Diversinervus paradisicus (Motschulsky)

Chiloneurus paradisicus Motschulsky (1863: 52).

Taxonomy: Compere (1931: 233-234); Trjapitzin (1961: 669-671).

Distribution: Sri Lanka.

Host: Coccidae: Saissetia coffeae (Walker).

Diversinervus scutatus Compere

Diversinervus scutatus Compere (1931: 235-237).

Distribution: South Africa.

Host: Coccidae: Ceronema sp.

Diversinervus silvestrii Waterston

Diversinervus silvestrii Waterston (1916: 137-140).

Distribution: Mauritius.

Host: Coccidae: Coccus viridis (Green).

Diversinervus smithi Compere

Diversinervus smithi Compere (1940: is

Biology: Flanders (1939, 1952).

Distribution: South Africa.. Released in California but failed to become

established (Bartlett 1978).

Host: Coccidae: Saissetia:dleae (Olivier).

Diversinervus stramineus Compere

Diversinervus’stramineus Compere (1938: 321-322).

Distribution: South Africa.

Host: Saissetia persimilis (Newstead).

SUBGENERIC CLASSIFICATION

The known members of Diversinervus seem to fall naturally into two

species groups:

The elegans group: Female with tufts of bristles on both the meso-

scutum and scutellum. In members with known males, the head of the male

is dark metallic. Species in this group have banded eggs, and first-instar lar-

vae with open tracheal systems. This group presently includes desantisi,

elegans, madgaoensis, paradisicus, scutatus, silvestrii and stramineus.

The smithi group: Female lacking a tuft of bristles on the mesoscutum.

In members with known males, the head of the male is yellow. Species in

this group have unbanded eggs, and first-instar larvae with closed tracheal

systems. This group presently includes cervantesi, masakaensis and smithi.

Although available information appears to support the establishment of

these tentative groups, their validity and relationships are presently unclear.

Rosen & Alon: Diversinervus (Encyrtidae) 359

For one thing, the males of masakaensis, paradisicus and silvestrii are

still unknown, the male of stramineus was described without a head, and im-

mature stages have not yet been described for most species. In the absence

of a mesoscutar tuft as well as in immature characters, the smithi group

appears to be more closely related to the genus Cheiloneurus than is the

elegans group. On the other hand, however, madgaoensis, a member of the

eleans group, resembles Cheiloneurus in having an elongate marginal vein in

the female fore wing. For the time being, therefore, establishment of inform-

al species groups appears preferable to formal classification into subgenera.

KEY TO THE SPECIES OF DIVERSINERVUS, FEMALES*

Lo > Mesoscutart tultoabsent (4. 9. oe ea ce Siw eye The smithi group. ... 2

NMesoscutat tum present. oA0 0°... 3. ee 2 The elegans group... . 4

2. Frontovertex about one-fourth width of head, deeply emarginate poster-

iorly; antennal scape moderately expanded, clearly more than twice as

long as wide; at least some funicular segments as long as wide, or

longer; club shorter than the funicle and not much wider; ovipositor

exserted somewhat; an arcuate, dark brown cross-band on face and

cheeks

Frontovertex wider, one-third width of head, not emarginate posteriorly;

scape strongly expanded, twice as long as wide; all funicular segments

wider than long; club large, as long as the funicle and considerably

wider; ovipositor concealed; face and cheeks without a dark cross-—band

bie iar BO 3 pee a PON: 5 EE Gee eee eee el ewe a ek 2 eae ae 2S

3. Wings greatly reduced; mesonotal and metanotal sclerites reduced, pro-

notum, tegulae and mesopleura relatively large; ocelli small, the

posterior pair 5 times their own diameter from occipital margin;

funicular segment V lined with dark brown dorsally; middle coxa dark

DIGWH sass 2G ; on ts eee eee — wn 4 Cerrantes

Wings fully developed; thoracic tergites normal; 6celti large, the posterior

pair twice their own diameter from occipital margin; funicular segment

NV. entirely yellow; middle coxa white). ooo) wie secs 6 oh ee BMIENT

4. Frontovertex densely reticulate-punctate, appearing granular ...... 5

Frontovertex smooth, shining, with tiny setigerous punctations ..... 6

5. Scape strongly expanded, twice as long as wide; marginal vein of fore

wing subequal in length to the stigmal vein; general coloration clay

VOUOW ee Ry Re es Oe Ba, ee ee ee ee

Scape moderately expanded, more than twice as long as wide; marginal

vein twice as long as the stigmal vein; general coloration dark

6. Scape moderately expanded, more than twice as long as wide .. x... 7

Scape strongly expanded, less than twice as long as wide; club large,

nearly as long as the funicle; mesoscutar and scutellar tufts short,

BO ee ie eR ee ee ee ms Ge eo mde Mae ore ep Us em elmer gel Ug ota Nig he mee mane Aa

*D. intermedius Hayat, Alam and Agarwal not included.

360 . Contrib. Amer. Ent. Inst., vol. 20, 1983

7. General coloration yellow; mesoscutum with a narrow metallic cross—band

CN. POSEETIOR MIO Lo testis nes ag ina BW Redeye lwody ease goer chins 8

General coloration extensively metallic; mesoscutum golden to brownish-

orange on either side of tuft, anterior part and sides black, with a

wide metallic cross~band on posterior one-half; scape nearly 3 times

as long as wide; club large, nearly as long as the funicle

8. Ocellar triangle acute; scape only a little more than twice as long as wide;

club distinctly shorter than the funicle; gaster subequal in length to

Cie nee ee a a ee ra Ue, Cae ohne tan Cee

Ocellar triangle right-angled; scape nearly 3 times as long as wide; club nearly

as long as the funicle; gaster considerably shorter than the thorax . .

Beg gles RN ast ete kat re aE Os et NS A bale ici ENS gS ge GN ge gS silvestril

9. Frontovertex very narrow, 6 times as long as wide, occupying one-ninth

width of head; ovipositor exserted to one-eighth length of gaster; fore

Wine Ofoed, "Twice as lene ae wie oe eee paradisicus

Frontovertex somewhat wider, about 4 times as long as wide, occupying

one-sixth to one-fifth width of head; ovipositor exserted to one-

fourteenth length of gaster; fore wing narrower, nearly 3 times as long

RI Ao An eal a TER Utes ak RaW i hE aN a GM takik Gy Gee aha. UR Seah a elegans

ACKNOWLEDGMENTS

The authors are grateful to Dr. John S. Noyes, British Museum (Natural

History), London, for his help in the identification of D. cervantesi and for

his comments on D. silvestrii, and to Dr. E. C. Dahms, Queensland Museum,

Brisbane, for loan of the types of cervantesi.

This research was supported in part by a grant from the United States -

Israel Binational Science Foundation (BSF), Jerusalem.

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Annecke, D. P. and Insley, H. P. 1971. Catalogue of Ethiopian Encyrtidae and

Aphelinidae (Hymenoptera: Chalcidoidea). Ent. Mem. Dept. Agric. Tech.

Serv. S. Afr. 232) 1-53.

Bartlett, B. R. 1962. The ingestion of dry sugars by adult entomophagous

insects and the use of this feeding habit for measuring the moisture needs

of parasites. J. Econ. Ent. 55: 749-753.

Bartlett, B. R. 1978. Coccidae. In: Clausen, C. P. (Editor), Introduced

parasites and predators of arthropod pests and weeds: a world review.

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Diversinervus elegans (Encyrtidae: Hymenoptera), an imported parasite

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976.

Rosen & Alon: Diversinervus (Encyrtidae) 361

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the British Museum. Bull. Ent. Res. 29: 315-337.

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but of Nature's bosom, notably new insects. Priv. Publ., Brisbane, 5 pp.

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encyrtid parasites (Hymenoptera: Encyrtidae) in India. Aligarh Musl.

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in Israel:. an annotated list. J. Ent. Soc. S. Afr. 43: 113-128.

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morphological adaptations for respiration. Univ. Calif. Publ. Ent. 8: 25-

ine.

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362 Contrib. Amer. Ent. Inst., vol. 20, 1983

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Hymenoptera). Canad. Ent. Suppl. 30: 1-1092.

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10th Ed. Edward Bros., Ann Arbor, Mich. 435 pp.

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Israel. Beitr. Ent. 17: 251-279.

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Res. 7: 137-144.

Prof. David Rosen is at present Cooperating Scientist, Systematic

Entomology Laboratory, IIBIII, Beltsville Agricultural Research Center,

USDA, Beltsville, MD 20705.

A TAXONOMIC STUDY OF THE SOUTH AMERICAN GENUS MEGALOPROCTUS

SCHULZ (HYMENOPTERA : BRACONIDAE : DORYCTINABE)

Paul M. Marsh

Systematic Entomology Laboratory, U.S.D.A.

Washington, D.C. 20560

The Neotropical Region is rich in species and genera of the braconid sub-

family Doryctinae. However, as is the case for most of the Braconidae, the

Doryctinae are largely unstudied and many genera and species are undescribed.

The South American genus Megaloproctus Schulz is one of the most

strikingly colored and largest in size of the Western Hemisphere Doryctinae.

The body length ranges from 8 to 20 mm and the ovipositor of some species

reaches 30 mm. This long ovipositor would indicate a parasitic habit on hosts

which bore deeply into wood or stems, but there are no biological records for

any species in the genus.

Fischer (1980) reviewed the genus based on ten specimens in the Hungarian

Natural History Museum, Budapest, and provided a key to seven species. In

the present study, based on more than 200 specimens, I have described 13

species, five of which are new, and prepared keys to females and males.

Most the specimens for this study are from the collections of the U-S.

National Museum of Natural History, Washington, D.C. and the British Museum

(Natural History), London. Other specimens, including holotypes of several

species, have been borrowed from the following collections: American Museum

of Natural History, New York; California Academy of Sciences, San Francisco;

Canadian National Collection, Ottawa; Carnegie Museum, Pittsburgh, Pennsylvania;

Cornell University, Ithaca, New York; Fundacion Miguel Lillo, Tucuman,

Argentina; Hungarian Natural History Museum, Budapest; Museum of Compara-

tive Zoology, Harvard University, Cambridge, Massachusetts; Natural History

Museum, Paris; Polish Academy of Sciences, Warsaw; Rijksmuseum van Natuur-

lijke Historie, Leiden; Swedish Museum of Natural History, Stockholm. I ex-

press my appreciation to the individuals from each of these institutions who

arranged for the loan of specimens.

Genus MEGALOPROCTUS Schulz

Megaproctus Brullé, 1846, p. 467. Preoccupied by Schoenherr, 1838.

Type-species: Megaproctus gt guius Brullé, designated by Viereck, 1914.

Megaloproctus Schulz, 1906, p. 138. Emendation.

Megistoproctus Schulz, 19ll, p. 88. New name for Megaproctus Brullé.

Prosthiacantha Enderlein, 1912, p. 24. Synonymy by Shenefelt and Marsh, 1976.

Type-species: Prosthiacantha harpactorina Enderlein. Monotypic.

Ectetamenochir Enderlein, 1912, p. 33. Synonymy by Roman, 1924.

Type -species: Ectetamenochir crinicornis Enderlein. Monotypic.

Schulz (1906) emended the name Megaproctus to Megaloproctus, but

there is no indication that he was aware that Megaproctus was preoccupied.

Later, Schulz (1911) proposed a new name, Megistoproctus, for Megaproctus

363

364 Contrib. Amer. Ent. Inst., vol. 20, 1983

apparently without realizing that he had changed the name earlier. According

to Articles 19 and 33a of the Code of Zoological Nomenclature, regardless of

whether Megaloproctus was a justified or unjustified emendation, it is an

available name.

Description: Moderate to large species, length of body 8-20 mn, |

length of ovipositor 5-30 mm; head cubical, occipital carina present, eyes

large, malar space 1/3 eye height or less, face ruguose (Figs. 20-24), rest of

head smooth, vertex occasionally strigate (Figs. 10, 12); thorax smooth, with

limited amount of sculpturing, notauli impressed and weakly scrobiculate, pro-

podeum nearly horizontal, without a distinct posterior declivity; abdomen long

and narrow, usually smooth, second tergum always with a smooth, arcuate,

transverse groove across middle; fore wing (Figs. 1-8) with three cubital

cells, recurrent vein entering first cubital cell or interstitial with first inter-

cubitus, first and second segments of mediella and basella of hind wing about

equal in length, rarely first segment of mediella longer than second; legs long

and slender, fore tibia with row of stout spines on anterior edge (Figs. 22,

26), hind coxa rounded ventrally at base, without a tooth; ovipositor at least

as long as abdomen, often longer than entire body.

Distribution: Central and South America.

Biology: Unknown. The extremely long ovipositor suggests that species

are possibly parasites of beetle larvae which bore deeply in wood.

Megaloproctus is related to Ontsira Cameron and Doryctes Haliday by

wing venation but is distinguished by the hind coxa being rounded ventrally at

the base and without a tooth. In habitus and coloration it is similar to

Binarea Brulle but is easily distinguished by the presence of the occipital

carina.

All terms for sculpturing are based on Harris, 1979.

KEY TO THE SPECIES

Females

1. Wings banded with alternate yellow and brown (Figs. 2, 4, 6, 8) ... 2

Wings not banded, completely infuscated, stigma occasionally yellow . .

Ny Gh UN OR Ce Meine aur Og eh ak Gi gh 7

2. Ocell-ocular distance greater than length of lateral ocellus and equal to

or greater than distance between lateral ocelli (Figs. 13, 14) ... .3

Ocell-ocular distance equal to or less than length of lateral ocellus and

always shorter than distance between lateral ocelli (Figs. 9, 10, 15) .

ee a ih ae ais cy a ae anime AON CMR Ua IIMS UR i a Ra

3. Ovipositor longer than entire body; fore tarsus more than two times

length Of Tore tibia es ee ee 2. brasiliensis (Szépligeti)

Ovipositor slightly longer than abdomen; fore tarsus less than two times

TRUE SE Berek CE e hen pad) + Oe ate Ea Ayasleibene hae ladoy buercaccue ee 4

4. Radiellen cell of hind wing with weak cross vein; hind femur and tibia

yellow; fore tibia with single row of stout spines along anterior edge

mB abil Pa Need ih, eben” Ss la a ell aU 7. bifasciatus (Szépligeti)

Marsh: Megaloproctus (Braconidae) 365

Radiellen cell without cross vein; hind femur and tibia black; fore tibia

with double row of 9-13 stout spines (Fig. 26) . 4. Pee ines (Brullé)

5. Radiellen cell of hind wing without weak cross vein (Fig. 2) ......

PROP Ry ME PO ere ONE EMA ISE Ne MARS 1 Ven elie, Ai get 6. harpactorinus (Enderlein)

Radiellen, cell. wath, weak :ctogs vei (PF ige.5 44.6) o imicnnsds cvelacaichinsy 9

6. First abdominal tergum, at least laterally, and base of second longitudi-

nally stricatentugone (Migs I) cae ea yl 10. striatus, MN. sp.

ADGGWHNAL TOTen. SINNOIN Ca ck ea ee ww kek 9. schunckei, N. Sp.

7. Body’ unicolorous: brown: stigma vyellow (Pigs. 1 3) 3s ew. ke 6

At least mesothorax differently colored than head; stigma black or brown

ae Goes eld eR ig eooe hegre Tae BAN a ea ek ihe ea 9

8. Vertex strigate between lateral ocelli (Fig. 12); apical antennal segments

beard) sierra oes a a gs 3. castaneus (Brullé)

Vertex smooth (Fig ll); apical antennal segments flattened .......

WT chacins hs aaa init he ps ORs fea ee iy ol aot iat) 2 Oe od eee eh aD.

9. Radiellen cell of hind wing divided by cross vein

MPP aRED Togo MEN Sa ye an aici Dana aie Sed enad § pecohaetl)

10. Abdominal terga 1 and 2 strigate-rugose; thorax, abdomen, and hind coxa

MEO PENANG Gis Pie He ee igs ha elas 13, xanbhas.’ 5... Mc Sp:

Abdominal terga | and 2 smooth; thorax and abdomen at least partly

DROS UG RR AIO ki ae eles Cha eee elas hh att ae il 11

11. . Abdomen entirely black; ovipositor longer than body; body length greater

Chan AS A a Rr a TTS et agg IZ: \townesi, ni sp.

Abdomen partly orange, ovipositor shorter than body; body length less

Freer Wi. Vi 5 Gi Ce Fe ey pee One ean A es ea 12

12. Radial cell of fore wing nearly five times as long as greatest width

(Fig. 5); radiellen cell of hind wing without curved infuscated spot

at base; fore tarsus about two times as long as fore tibia .....

ae Me LL gk Fe ae Ns OME elles on et ae ae 11. strongylogaster (Cameron)

Radial cell not more than four times as long as width (Fig. 7);

radiellen cell with curved infuscated spot at base; fore tarsus about

l-lj2 times ag: tong .agvtore tibia oi. uss 7. pertinax (Cameron)

Males

The following key is not very reliable. As is often the case in the

Braconidae, males are difficult to identify unless associated in rearings with

females. I have seen no associated sexes and only a few male specimens, so

the key is based on males that come closest to agreeing with the females.

1. Wings not banded, uniformly infuscated ......... ig Vids tt ee, Sa 2

Wings Bates We. OP FAO Re i hae: Rah i ale abs Y i aoe

Ae . OGY MAACO IGE OIS TNEGNE OUTS ae NN 3

Body yellow of oranee and black, stigma black o.oo a Uk 4

3. Vertex strigate between lateral ocelli. ...... 3. castaneus (Brullé)

ETON MONG es a ee ieee hit ce i ech 8. platyantennus, nN. sp.

366 Contrib. Amer. Ents Inst/,ovele 20; 1983

4. Thorax and basal abdominal segments orange; head, most of legs, and

apex or ubdomen black 20° 2 re 5. Pt pennies (Szépligeti)

Body enewrely yeliow u4i5 0. Ge ey ee 7. pertinax (Cameron)

5. Ocell-ocular distance less dus length of lateral ocellus ........ 6

Ocell-ocular distance equal to or greater than length of lateral ocellus

Oe aes Be ae Bee a ea a ee es es Haga a |

6... Vertex Siig0th . 248.6 inl ete ee 10° striatus, nN. sp-

Vertex striate oc poct5 ghee Ate 6. harpactorinus (Enderlein)

7. Fore tarsus more than two times as long as fore tibia.........

ee 2. brasiliensis _(Szépligeti)

Fore tarsus less than two times as long as fore tibia. .........

Cole ie ec STE Ho a a dadumus (Braille)

1. Megaloproctus bifasciatus (Szépligeti)

Megaproctus bifasciatus Szépligeti, 1904, p. 192. Holotype female in

Hungarian Natural History Museum, Budapest.

Megaloproctus bifasciatus (Szépligeti); Shenefelt and Marsh, 1976, p.

1319.

Female: Length of body, 12 mm; ovipositor 10 mm. Color: head black,

palip yellow, mouth area orange; thorax black, upper part of propleuron,

mesopleuron above subalar groove, tegulae and mesonotal lobes orange; ab-

dominal terga 1-5 orange, terga 6-7 black, terga 4 and 5 with black lateral

spots; fore leg yellow except last tarsomere black, mid leg yellow except coxa

brown and last tarsomere black, hind leg with coxa, trochanter |, base of

trochanger 2, apex of femur, base and apex of tibia, and last 4 tarsomeres

black, apex of trochanter 2, rest of femur and tibia, and first tarsomere

yellow; wings with two dark cross bands (similar to Aiaunns). Head: vertex,

frons, and temples smooth, face rugose, rugae running transversely at sides;

ocell-ocular distance about two times as long as lateral ocellus; frons slight-

ly depressed, with longitudinal shallow furrow; first flagellomere equal in

length to second; longest segment of maxillary palpus equal in length to first

flagellomere. Thorax: Pronotum and proepisternum with deep hair sockets

giving punctate appearance; subalar groove weakly scrobiculate, sternaulus and

mesopleural furrows scrobiculate; propodeum smooth, hair sockets deep, med-

ian basal carina long and distinct, areola not margined, short longitudinal

carina at base above each spiracle. Abdomen: smooth, first tergum slightly

longer than apical width; ovipositor slightly shorter than body. Legs: fore

tibia with row of 5 stout spines along anterior edge and 6 spines at apical

edge; fore tarsus about 1-1/2 times as long as fore tibia. Wings: recurrent

vein of fore wing interstitial with first intercubitus; second segment of radius

short, less than two times as long as first segment; nervulus postfurcal by

distance greater than its length; radiellen cell of hind wing widening toward

apex, with weak cross vein at basal 1/3; first and second segments of

mediella about equal in length. (The fore wings each have a vein in the anal

cell near the nervulus which angles toward the wing apex. This is not a

characteristic vein even for the subfamily and I consider it to be an anomaly.)

Male: Unknown.

Marsh: Megaloproctus (Braconidae) 367

Type locality: Marcapata, Peru.

Distribution: Peru. Known only from the holotype.

This species is very similar to didymus and schunckei. From didymus it

is separated by the cross vein in the radiellen cell of the hind wing and the

yellow hind legs; from schunckei it is distinguished by the greater ocell-

ocular distance.

2. Megaloproctus brasiliensis (Szépligeti) (Figs. 8, 14, 19, 22)

Megaproctus brasiliensis Szépligeti, 1902, p. 59. Holotype female in

Hungarian Natural History Museum, Budapest.

Megaproctus peruiensis Szépligeti, 1902, p. 60. Holotype female in

Hungarian Natural History Museum, Budapest.

Ectetamenochir crinicornis Enderlein, 1912, p. 34. Holotype female in

Polish Academy of Sciences, Warsaw.

Megaloproctus brasiliensis (Szepligeti); Shenefelt and Marsh, 1976, p.

1319.

Female: Length of body, 11-16 mm; ovipositor, 14-25 mm. Color: head

and antenna black; thorax varies from entirely orange to entirely black, more

often orange with proepisternum black; all coxae, hind trochanter | and hind

femur black, rest of fore and middle legs yellow except femora which are

occasionally light brown, hind tibia and tarsus dark brown to black, apex of

hind femur and base of hind tibia occasionally dark yellow, hind coxa oc-

casionally orange on basal 1/3; abdominal terga 1-4 always orange, tergum 7

always black, terga 5-6 varying from black to orange; wings banded but some-

times not distinctly. Head: face finely rugose (Fig. 22), vertex and temples

smooth, frons slightly depressed and occasionally rugose and with a scrobicu~

late median longitudinal groove (Fig. 14); ocell-ocular distance two times

length of lateral ocellus; antenna 80-86 segmented, first flagellomere about

as long as second; longest segment of maxillary palpus about equal in length

to second flagellomere. Thorax: propodeum (Fig. 19) with median basal

carina usually distinct, areola indistinctly defined by rugosities. Abdomen

(Fig. 19): entirely smooth; basal corners of second tergum depressed; first

tergum slightly longer than apical width; ovipositor longer than entire body,

1-1/4 to 1-1/2 times longer. Legs: fore tibia with 4-5 spines along anterior

edge, 8-9 spines at apical edge: fore tarsus 2-1/4 times as long as fore

tibia. Wings (Fig. 8): recurrent vein entering first cubital cell; nervulus post-

furcal by nearly its own length; first segment of mediella usually shorter than

second segment.

Male: Essentially as in female except hind femur orange and body length

shorter.

Type localities: Massanary, Brazil (brasiliensis); Pebas, Peru

(peruiensis); Obidos, Brazil (crinicornis).

Distribution: Venezuela: Mt. Marahuaca. British Guiana: Mazarumi,

Kartobo, Tumatumari. Surinam: Para District, Mooi Wane, Zanderij, Mapane,

Albina~Moengo, Coppename Bakhius Mts., Poeloegoedoe. French Guiana: Mana

River. Peru: Pucallpa, Pebas. Brazil: Massanary, Obidos, Para, Manaos, Rio

Autaz.

368 Contrib. Amer. Ent. Inst., vol. 20, 1983

This species is easily distinguished from the other species with banded

wings by its long ovipositor being distinctly longer than entire body.

3. Megaloproctus castaneus (Brullé) (Figs 1, 12, 21)

Megaproctus castaneus Brullé, 1846, p. 468. Holotype female in Natural

History Museum, Paris.

Megaproctus xanthostigma Szepligeti, 1904, p. 192. Holotype female in

Hungarian Natural History Museum, Budapest.

Bracon? ferruginosus Holmgren, 1869 (1868), p. 426. Synonymy by Roman,

1910. Holotype female in Swedish Natural History Museum, Stockholm.

Megaloproctus castaneus (Brullé); Shenefelt and Marsh, 1976, p. 1319.

Female: Length of body, 10-19 mm; ovipositor, 12-20 mm. Color: entire

body red-brown except fore and mid legs which are honey yellow; wings

dusky brown, stigma and small area below bright yellow. Head: face (Fig.

21) rugose with definite transverse rugae on each side, vertex strigate between

lateral ocelli and with a sharp transverse ridge between lateral ocelli so that

frons and vertex are nearly at a right angle to each other (Fig. 12); ocell-

ocular distance about equal to length of lateral ocellus; antenna 70-75 seg-

mented, first flagellomere shorter and thicker than second, all flagellomeres

longer than wide, apical ones not flattened as in platyantennus; longest seg-

ment of maxillary palpus a little longer than second flagellomere. Thorax:

propodeum smooth, without stub of median basal carina. Abdomen: first

tergum about as long as apical width, smooth dorsally except for apical later-

al corners which are occasionally rugose; terga 2~3 smooth except for some

rugosities at base, second tergum with weak converging lines from dorsal cor-

ners to suturiform articulation; rest of terga smooth; ovipositor about as long

as body, sharply curved downward at tip. Legs: fore tibia with 5-7 spines

along anterior edge, 7-8 spines at apical edge; fore tarsus about two times

as long as fore tibia; hind femur about 3-l/2 times as long as wide, widest

near middle. Wings (Fig. 1): second segment of radius longer than second

intercubitus, first cubital cell not as short as in platyantennus; recurrent vein

entering first cubital cell; nervulus postfurcal by more than its length; first

and second segments of mediella and basella about equal in length to each

other; radiellen cell noticeably widening at apex.

Male: Essentially as in female.

Type localities: Rio Grande Province, Brazil (castaneus); Rio de

Janeiro, Brazil (ferruginosus); Paraguay (xanthostigma).

Distribution: Brazil: Rondon, Cauna, Nova Teutonia, Rio de Janeiro,

Blumenau, Rio Grande Province. Paraguay: no locality. Argentina: Tucuman.

This species is similar to platyantennus in habitus and color; it can be

easily distinguished from platyantennus by its normal un-flattened apical an-

tennal segments and strigate vertex.

4. Megaloproctus didymus (Brullé) (Figs. 13, 16, 20, 26)

Megaproctus didymus Brullé, 1846, p. 467. Holotype female in Natural

History Museum, Paris.

Marsh: Megaloproctus (Braconidae) 369

Megaproctus affinis Szépligeti, 1904, p. 192. New synonymy. Holotype

female in Hungarian Natural History Museum, Budapest.

Megaproctus unifasciatus Szépligeti, 1906, p. 601. New synonymy. Holo-

type female in Hungarian Natural History Museum, Budapest.

Megistoproctus marcapatanus Roman, 1924, p. 22. New synonymy. Holo-

type female in Swedish Natural History Museum, Stockholm.

Megistoproctus autazicus Roman, 1924, p. 23. New synonymy. Lecto-=

type female in Swedish Natural History Museum, Stockholm.

Megaloproctus didymus (Brullé); Shenefelt and Marsh, 1976, p. 1319.

Female: Length of body, ll1-13 mm; ovipositor, 9-11 mm. Color: head

and antenna black, clypeus dark red; prothorax anterior to propleural groove,

apex of scutellum and propodeum black, mesothorax and rest of prothorax

orange, mesopleuron rarely with black markings; first and second abdominal

terga and base of third orange, rest of terga black; coxae black, trochanter

1 black or yellow, trochanter 2 always yellow, fore femur, tibia and tarsus

yellow, mid femur usually yellow, rarely black, mid tibia and tarsus yellow,

hind femur, tibia and tarsus black. Head: vertex and temples smooth, face

coarsely rugose (Fig. 20), frons smooth, slightly depressed and with shallow

median scrobiculate groove (Fig. 13); ocell-ocular distance about 1-1/4 times

length of lateral ocellus; antenna 67-74 segmented, first flagellomere slightly

longer than second. Thorax: propodeum with distinct strong basal carina and

strongly margined areola (Fig. 16); subalar groove usually scrobiculate. Abdo-

men: entirely smooth; first tergum slightly longer than apicai width; second

tergum with basal corners depressed; ovipositor shorter than body, slightly

longer than abdomen. Legs: Fore tibia with scattered row of 9-13 spines

on anterior edge, 8-9 spines at apical edge (Fig. 26); fore tarsus about 1-3/5

times longer than fore tibia, sometimes longer. Wings: recurrent vein enter-

ing first cubital cell; nervulus postfurcal by about its own length; first seg-

ment of mediella of hind wing slightly shorter than second.

Male: Essentially as in female; fore tibia with single row of 6-8 spines.

Type localities: la Guyane (probably French Guiana) (didymus);

Marcapata, Peru (affinis and marcapatanus); Mapiri, Bolivia (unifasci-

atus); Rio Autaz, Brazil (autazicus).

Distribution: Trinidad: New York Zoological Society Tropical Research

Station. British Guiana: Kartabo, Essequibo River, Demerara River. French

Guiana: Mana River. Peru: Marcapata, Yanayacu, Boqueron Abad. Bolivia:

Beni, Tumupasa, Mapiri. Brazil: Santarem, Apipica, Coraci, Igarape Gurupi,

Campinas, Feija.

This species is similar to brasiliensis but distinguished by its shorter

ovipositor (shorter than body), shorter fore tarsus (less than twice length of

fore tibia), relative lengths of segments of the mediella, and its coarsely

rugose face. There is considerable variation in color in the specimens that

I have seen with three different types following certain geographical bound-

aries. However, in view of the variation found in other species, such as

harpactorinus, and until I have seen more specimens, | prefer not to place

these groups into subspecies.

370 Contrib. Amer. Ent. Inst., vol. 20, 1983

Roman described autazicus from one female and four males but did not

designate a holotype. I have seen the female and two males from this series

and hereby designate the female as lectotype.

5. Megaloproctus fumipennis (Szépligeti)

Megaproctus fumipennis Szépligeti, 1904, p. 191. Holotype female in

Hungarian Natural History Museum, Budapest.

Megaloproctus fumipennis (Szépligeti); Shenefelt and Marsh, 1976, 1319.

Female: Length of body, 6.5 mm; ovipositor, 4.5 mm. Color: clypeus and

malar space orange, rest of head black; antenna black; pro- and mesothorax

orange; propodeum black; abdominal terga 1-3 orange, 4-8 black; fore and mid

legs orange except base of tibiae black, hind leg black except apex of second

trochanter and apex of femur orange; wings uniformly brown. Head: vertex,

frons and temples smooth; ocellocular distance shorter than diameter of lateral

ocellus; eyes large, temples less than half width of eye; face rugulose; anten-

nae of type broken. Thorax: propodeum with short basal median carina, long

narrow pentagonal areola, and lateral longitudinal carinae above each spiracle;

rugose inside areola and around spiracles. Abdomen: first tergum as long as

apical width, very weakly strigate; ovipositor shorter than body. Legs: fore

tibia with 8 spines on anterior edge and 5 spines on apical edge; fore tarsus

1-1/3 times as long as fore tibia. Wings: recurrent vein entering first cubital

cell at apex; nervulus postfurcal by slightly less than its length; radiellen cell

widening toward apex, divided in basal 1/3 by weak cross vein; first segment

of mediella slightly longer than second segment.

Male: Essentially as in female; propodeum orange, proepisternum black.

Type locality: Sicuani, Peru.

Distribution: Peru. In addition to the holotype, I have seen only two

males from Peru that are apparently this species. They are labeled as being

from Chauchamayo which I could not locate. This locality is probably a mis-

print for Chauchosmayo River which is very near the type locality of Sicuani.

The distinguishing features of fumipennis are its color, sculpture on pro-

podeum, and cross vein in the radiellen cell of the hind wing.

6. Megaloproctus harpactorinus (Enderlein) (Figs. 2, 10, 23, 25)

Prosthiacantha harpactorinus Enderlein, 1912, p. 25. Holotype male in

Polish Academy of Science, Warsaw.

Megaproctus nigridorsum Enderlein, 1920, p. 137. Holotype female in

Polish Academy of Science, Warsaw.

Megistoproctus debilis Roman, 1924, p. 21. Holotype female in British

Museum (Natural History), London.

Megistoproctus opaculus Roman, 1924, p. 24. Holotype female in

Swedish Natural History Museum, Stockholm.

Megaloproctus harpactorinus (Enderlein); Shenefelt and Marsh, 1976,

Poloe0.

Female: Length of body, 10-17 mm; ovipositor, 10-20 mm. Color: head

black, malar space and clypeus occasionally orange; antenna black; thorax

Marsh: Megaloproctus (Braconidae) 371

varying from entirely orange to entirely black; abdomenal terga |-3 always

orange, terga 7-8 always black, terga 4-6 varying from black to orange;

fore and mid legs always yellow except last tarsomere always and coxae oc-

casionally black; hind leg varys from entirely black to yellow except femur

always yellow at least at base; wings banded yellow anddark brown, apical

half of hind wing occasionally entirely infuscated. Head: vertex strigate be~

tween lateral ocelli (Fig. 10), with a transverse ridge between ocelli so that

vertex and frons are nearly at right angles to each other; lateral ocelli nearer

eyes than to each other, ocellocular distance about 3/4 diameter of lateral

ocellus face rugose with definite transverse rugae on each side (Fig. 23), an-

tenna 73-80 segmented, first flagellomere shorter than second; longest seg=

ment of maxillary palpus about equal to second flagellomere. Thorax: pro-

podeum with weak longitudinal median carina, areola absent, occasionally area

of areola rugulose. Abdomen: entirely smooth; apical corners of first tergum

and basal corners of second tergum impressed and occasionally rugulose; first

tergum longer than apical width; ovipositor slightly longer than body, curved

down at apex. Legs: fore tibia with 5-6 spines on anterior edge, 6-8 on

apical edge (Fig. 25); fore tarsus about 1-4/5 times as long as fore tibia; hind

femur about 3-3/4 times as long as wide, widest in middle. Wings (Fig. 2):

recurrent vein entering extreme apex of first cubital cell, occasionally appar-

ently interstitial with first intercubitus; nervulus postfurcal by at least its own

length, occasionally more than its length; radiellen cell gradually widening

toward wing apex.

Male: Essentially as in female.

Type localities: Obidos, Brazil (harpactorinus); Mucury, Brazil

(nigridorsum); Brazil, probably Amazonas (debilis); Rio Autaz, Brazil

(opaculus).

Distribution: Brazil: Rio caiary-Uaupes, Santarem, Rio Autaz, Obidos,

Mucury. Peru: Pucallpa, Chauchamayo, Huanaco, Yanayacu, Colonia Perene,

El Campamiento. British Guiana: Essequibo River. French Guiana: Mana

River. Bolivia: Prov. del Sara, Cuatro Ojos, Santa Cruz, Cochabamba.

Venezuela: El Tucuco.

This species is quite variable in color and similar in habitus to the other

species with banded wings, but it is easily distinguished by the strigate ver-

tex and short ocellocular distance (3/4 diameter at lateral ocellus).

7. Megaloproctus pertinax (Cameron), revised status (Fig. 7)

Doryctes pertinax Cameron, 1887, p. 383. Holotype female in British

Museum (Natural History), London. Incorrectly synonymized under

Megaloproctus strongylogaster (Cameron) by Shenefelt and Marsh,

1976, p. 1320.

Megaproctus fuscipennis Szépligeti, 1904, p. 191. Holotype female in

Hungarian Natural History Museum, Budapest.

Megistoproctus joachimi Roman, 1924, p. 19. Lectotype female in

Swedish Natural History Museum, Stockholm.

Female: Length of body, § 12 mm; ovipositor, 7-10 mm. Color: head

and antenna black; prothorax usually black with upper part of propleuron al-

ways orange, occasionally entire prothorax orange; mesothorax orange;

a2 Contribs Amer. Ent) inst: vole ZO, 1983

propodeum usually black, occasionally orange; abdominal segments 1-3 always

orange, segments 7-8 always black, segments 4-6 usually black but occa-

sionally orange; fore and mid tarsi yellow or light brown, apical tarsomeres

black; remainder of fore and mid legs and all of hind legs black; second

trochanter of all legs orange at apex; wings dusky black, usually with clear

spot in middle of first cubital cell and at junction of first intercubitus,

cubitus, and recurrent veins; radiellen cell with curved infuscated spot at

base. Head: vertex, frons, and temples smooth, face strongly rugose; teeth

of mandibles small, not crossing when closed; ocellocular distance about 1-1/2

times as long as diameter of lateral ocellus; frons slightly depressed, with

shallow scrobiculate groove between antennae; first flagellomere longer than

second; longest segment of maxillary palpus slightly longer than second

flagellomere. Thorax: propodeum smooth except occasionally rugose inside

areola and around spiracles, basal median carina and areola not always dis-

tinct. Abdomen: first tergum longer than apical width; second tergum with

basal corners slightly impressed; third tergum with smooth sinuate transverse

groove; entire abdomen smooth dorsally, rarely basal corners of second tergum

weakly rugose; ovipositor slightly shorter than body, slightly curved down at

apex. Legs: fore tibia with irregular row of 6-9 spines along anterior edge,

7 spines at apical edge; fore tarsi 1-1/2 to 1-2/3 times as long as fore tibia;

hind femur about 3-1/4 times as long as wide, widest in middle. Wings (Fig.

7): recurrent vein entering extreme apex of first cubital cell, rarely inter-

stitial with first intercubitus; nervulus postfurcal by slightly less than its

length; greatest length of radial cell about 3-3/4 times its greatest width;

radiellen straight to wing margin; radiellen cell with crescent shaped infus-

cated spot at base.

Male: Similar to female except color of body entirely orange with an-

tennae, tip of abdomen, hind trochanters, and hind tibiae and tarsi brown.

Type localities: Volcan de Chiriqui, Panama (pertinax, according to

label; description states Bugaba); Marcapata, Peru (fuscipennis); Rio Autaz,

Brazil (joachimi). Roman described joachimi from 53 specimens (44 M, 9

F, but did not designate a holotype in his paper although one of the syn-

types is labeled, apparently by him, as the holotype. I have selected this

specimen as the lectotype. The specimen is a female with the following data:

"Rio Autaz," "Amazon Roman," "66," "Typus" (red label), "Megistoproctus

Joachimi Rom. n.sp. F typ.," "245/70," "290/74," "Riksmuseum Stockholm,"

"Lectotype Megistoproctus joachimi Roman by P. Marsh 1974."

Distribution: Panama: Volcan de Chriqui (or Bugaba), Barro Colorado

Island. French Guiana: Mana River. Peru: Chauchamayo, Puerto

Maldonado, Pucallpa, Marcapata, Yanayacu, Boqueron Abad. Brazil: Rio

Autaz, Rio Purus, Benjamin Constant.

This species is similar to fumipennis but is distinguished by its black

legs, smooth propodeum, and lack of cross vein in radiellen cell of hind wing.

The infuscated spot in the radiellen cell may appear to be a cross vein but

is definitely not associated with any veins in the hind wing. This species

also can be confused with strongylogaster but is distinguished by the short-

er radial cell (less than 4 times its width), smaller size, shorter fore tarsus

(less than twice length of fore tarsus) and infuscated spot in the radiellen ,

cell of the hind wing.

Marsh: Megaloproctus (Braconidae) 373

8. Megaloproctus platyantennus, n. sp. ( Figs. 3, 11, 18, 24)

Female: Length of body, 8-14 mm; ovipositor, 8-14 mm. Color: head,

thorax, coxae, and abdomen reddish brown; legs honey yellow; antennae brown

at base, honey yellowin middle, black apically; wings uniformly dusky, light

brown, stigma and small area below bright yellow. Head: face rugose vertex

(Fig. 11) smooth between ocelli; lateral ocelli nearer to each other than to

eyes, ocellocular distance about 3 times diameter of lateral ocellus; antenna

65-70 segmented, first flagellomere about as long as second, flagellomeres

1 to about 30 longer than wide and not flattened, those from about 30-40

suddenly becoming shorter and flattened, those from 40 to apex of antenna

wider than long and very flattened; longest segment of maxillary palpus short-

er than second flagellomere. Thorax: pronotum with transverse carina across

apex extending onto propleuron; propodeum smooth with short stub of dorsal

median carina at base. Abdomen (Fig. 18): first tergum slightly longer than

wide, smooth dorsally except for rugose apical lateral corners; terga 2 and

3 with two scrobiculate longitudinal converging furrows and one usually

scrobiculate transverse sinuate furrow which set off raised u-shaped median

area at base;rest terga smooth; ovipositor about as long as body, straight or

only slightly curved down at apex. Legs: fore tibia with 5-7 spines along

anterior edge and 5 spines at apical edge; fore tarsus about 2-1/2 times as

long as fore tibia; hind femur about 4-1/3 times as long as wide, widest near

apex. Wings (Fig. 3): first cubital cell short, second segment of radius not

longer than second intercubitus; recurrent vein interstitial with first inter-

cubitus; nervulus postfurcal by less than 1/2 its length; second segment of

mediella slightly longer than first segment and basella; radiellen cell not

noticeably widening at apex.

Male: Essentially as in female; body length 7-10 mm; apical flagel-

lomeres not flattened.

Holotype: Female, Nova Teutonia, Brazil (27° 11'B, 52° 23'L), 7. Il.

1936, Fritz Plaumann. Deposited in the British Museum (Natural History),

London. Paratypes: Brazil: same data as holotype, dates from September

to February, 1937 to 1947, 39 F, 30 M; Rondon (24° 38'B, 54°07'L), 1 F,

7-IX-52, Fritz Plauman; Passe des Indies (27° 07'B, 52° 36'L), 1 F, 1 M, Il-

1937, F. Plaumann; Parana, 1 F, 1907-12, E. Dukinfield Jones. Paratypes de-

posited in U.S. National Museum,British Museum (Natural History), Canadian

National Collection and Swedish Museum of Natural History.

This species is similar to castaneus in habitus and color but can be

readily distinguished by the flattened apical segments of the female antenna,

the smooth vertex, and sculpturing of the abdomen. The flattened flagel-

lomeres of platyantennus are unique in this genus and, for that matter, in

the whole subfamily. I have seen one specimen identified as castaneus by

Roman which is actually platyantennus..

9. Megaloproctus schunckei , n. sp. (Figs. 6, 15)

Female: Length of body, 8-ll mm; ovipositor, 7-9 mm. Color: head

and antenna black, prothorax black except upper part of propleuron which is

orange, mesothorax orange; propodeum usually black, occasionally orange dor-

sally; abdominal terga 1-6 orange, tergum 7 black, terga 5-6 sometimes

marked with black at apex; fore legs yellow, fore coxae sometimes brown;

374 Contrib. Amer. Ent. Inst., vol. 20, 1983

mid leg yellow except coxa which are black; hind coxa and trochanters

black, hind femur yellow, hind tibia usually black, sometimes yellow on basal

half, hind tarsus black; wings banded. Head: vertex and temples smooth

(Fig. 15), face coarsely rugose, transversely so at sides, frons depressed,

smooth, with shallow longitudinal line between antennae; ocellocular distance

less than diameter of lateral ocellus; eyes large, malar space about 1/5 eye

height, temples less than 1/2 eye width; antennae broken in all specimens,

first and second flagellomeres about equal in length. Thorax: carinae on

propodeum weak, sometimes obscured, areola often a raised rugose area,

costulae absent. Abdomen: entirely smooth, first tergum longer than apical

width; ovipositor slightly shorter than body, curved down at apex. Legs: fore

tibia with row of 5-8 spines along anterior edge, 5-6 spines at apical edge;

fore tarsus 1-1/2 times as long as fore tibia. Wings (Fig. 6): recurrent vein

nearly interstitial with first intercubitus; nervulus postfurcal by two times its

length; first segment of mediella longer than second segment; radiellen cell

widening toward wing apex, with weak cross vein across basal 1/3.

Male: Unknown.

Holotype: Female, Pucallpa, Dept. Loreto, Peru, 29. IV. 1952, J. M.

Schuncke. Deposited in British Museum (Natural History), London.

Paratypes: 2 F, same data as holotype except dates of 8. V. 1952. De-

posited in U.S. National Museum.

This species is similar to striatus but differs in the smooth abdomen and

the weaker carinae on the propodeum.

10. Megaloproctus striatus , n. sp. (Figs. 4, 9, 17)

Female: Length of body, 8-ll mm; ovipositor, 6-9 mm. Color: head and

antennae black; pro- and mesothorax black, propodeum black or orange; ab-

dominal terga 1-6 orange, tergum 7 at least black on apical half; fore and

mid legs yellow, mid coxa sometimes brown, hind coxa black, hind trochan-

ters, femora and basal half of tibia yellow, apical half of tibia and tarsus

brown or black; wings banded. Head: vertex and temples smooth (Fig. 9),

face coarsely rugose, frons depressed, smooth; ocellocular line shorter than

diameter of lateral ocellus; eyes large, malar space about 1/6 eye height,

temples less than 1/2 eye width; antennae broken in all specimens, first and

second flagellomeres equal in length. Thorax: basal carina of propodeum

(Fig. 17) distinct but often obscured by rugosities, areola margined and ru-

gose, costulae present, second longitudinal carina present above spiracles.

Abdomen (Fig. 17): first tergum strigate-rugose laterally, basal half of sec-

ond tergum longitudinally strigate, rest of terga smooth; first tergum longer

than wide; ovipositor slightly shorter than body, curved down at apex. Legs:

fore tibia with row of 5-7 spines along anterior edge, 5-7 at apical edge;

fore tarsus about 1-1/3 times as long as fore tibia. Wings (Fig. 4): recur-

rent vein nearly interstitial with first intercubitus; nervulus postfurcal by

twice its length; radiellen cell widening toward wing apex, with a weak cross

vein at basal 1/3; first segment of mediellen longer than second segment.

Male: Essentially as in female; thorax entirely orange except prothorax

black.

Holotype: Female. Braganza, Para, Brazil, H. B. Merrill colr. Deposit-

ed in U.S. National Museum, Washington. Paratypes: Paraguay:

Marsh: Megaloproctus (Braconidae) 375

San Bernardino, | F, K. Fiebrig. Brazil: Corumba, 2 F. Venezuela, | F,

Paraitepuy, Bolivar, P. J. Andaze, Dec. 1940. Peru, | F, 1 M, Loreto,

Pucallpa, 13-VII-1951 and 24-1-1952, J. M..Schuncke. Deposited in U.S. Nation-

al Museum, British Museum (Natural History), Canadian National Collection,

and Cornell University.

This species is similar to bifasciatus and schunckei, but can be dis—

tinguished from bifasciatus by the shorter ocellocular distance and strigate

abdominal terga, and from schunckei by the strigate abdominal terga. It is

also the only species in the genus which has the costulae on the propodeum

clearly indicated. The two specimens from Brazil have a cross vein in the

anal cell of the fore wing similar to the holotype of bifasciatus but I con-

sider this also to be an anomaly.

11. Megaloproctus strongylogaster (Cameron) (Fig. 5)

Doryctes strongylogaster Cameron, 1887, p. 382. Holotype female in

British Museum (Natural History), London.

Megaloproctus strongylogaster (Cameron); Shenefelt and Marsh, 1976,

p. 1320.

Female: Length of body, 12-14 mm; ovipositor, 11-14 mm. Color: upper

part of propleuron, mesothorax, abdominal segments 1-3 and apex of second

trochanter orange; propodeum orange or black; rest of body and legs black;

wings evenly infuscated, not banded. Head: vertex, frons, and temples

smooth; ocellocular distance about 1-2/3 times diameter of lateral ocellus;

frons slightly depressed, with shallowscrobiculate groove between antennae; ~

face rugose, transversely so at sides; teeth of mandible large, crossing when

closed, second tooth large and truncate, longest segment of maxillary palpus

longer than second flagellomere. Thorax: subalar groove scrobiculate; pro-

podeum smooth except weakly rugose inside areola, basal median carina

strong, areola distinctly margined. Abdomen: mostly smooth, first tergum

and basal corners of second tergum weakly strigate in holotype; first tergum

longer than apical width; apical corners of first tergum and basal corners of

second tergum depressed; second tergum with transverse groove across mid-

dle; ovipositor about as long as body, slightly curved down at apex. Legs:

fore tibia with scattered row of 7-9 spines along anterior edge, 7 spines at

apical edge; fore tarsus about twice as long as fore tibia; hind femur about

3-1/2 times as long as wide. Wings (Fig. 5): recurrent vein entering first

cubital cell; nervulus postfurcal by slightly more than its length; length of

radial cell nearly five times its greatest width; radiellen vein straight to wing

margin, radiellen cell uniformly infuscated.

Male: Unknown.

Type locality: Bugaba, Panama.

Distribution: Brazil: Coraci. Peru: Rio Santiago, Surinam: Mapane.

This species is very similar to pertinax but can be separated by its

longer radial cell (nearly 5 times width) and longer fore tarsus (about 2

times length of fore tibia). From fumipennis it can be distinguished by the

absence of a cross vein in the radiellen cell of the hind wing.

376 Contribst Amer: Enty Inst.; vol. 20, °° 1983

12. Megaloproctus townesi, Nn. sp-

Female: Length of body, 16-19 mm; ovipositor, 22-28 mm. Color: body

black except propleuron, entire mesothorax, basal abdominal sterna, lateral

longitudinal lines on abdominal tergum 2, and apex of second trochanter which’

are bright orange; wings very dark brown with clear spots in center of first

cubital cell and at junction of cubital, first intercubitus, and recurrent veins.

Head: somewhat transverse, vertex in anterior view considerably higher than

upper level of eyes; antennae broken, first flagellomere longer than second;

longest segment of maxillary palpus longer than second flagellomere; vertex

smooth; ocellocular distance about twice as longas diameter of lateral ocellus.

Thorax: propodeum with strong basal median carina and two short lateral

basal carinae, areola not margined by carinae but indicated by pattern of

rugae. Abdomen: first tergum longer than apical width, smooth except few

rugae at base and apical corners; second tergum smooth with basal lateral

corners impressed and rugose; remainder of terga smooth; ovipositor longer

than body. Legs: Fore tibia with double row of 8-12 spines on anterior edge

and 7 spines on apical edge; fore tarsus 2-1/2 times as long as fore tibia;

hind femur 4 times as long as wide, widest in middle. Wings: recurrent

vein entering extreme apex of first cubital cell; nervulus postfurcal by more

than its length; radiella suddenly curved down at apex.

Male: Unknown.

Holotype: Female, Pucallpa, Loreto, Peru, Feb. 20, 1962, J. M. Schunke.

Deposited in British Museum (Natural History), London. Paratype: One fe-

male with same data as holotype but dated Dec. 21, 1959. Deposited in U.S.

National Museum.

This striking species is easily recognized by its large size, long ovipositor

(nearly 1-1/2 times longer than body), and color which is largely black. |

take great pleasure in naming this species after my good friend and col-

league, Henry Townes.

13. Megaloproctus xanthus , n. sp.

Female: Length of body, 10 mm; ovipositor, 7 mm. Color: head and

antenna black; proepisternum black, rest of thorax yellow orange; abdomen

yellow orange; fore legs black except apex of second trochanter and tarso-

meres 1-4 yellow orange, mid leg black except apex of second trochanter

and base of basitarsomere yellow orange; hind legs black except coxae, apex

of second trochanter and base of basitarsomere yellow orange; wings dusky

brownish yellow. Head: vertex smooth; ocellocular distance 1-3/4 times as

long as diameter of lateral ocellus; antenna 65 segmented, first flagellomere

equal in length to second; longest segment of maxillary palpus longer than

second flagellomere. Thorax: propodeum with short median basal carina,

long narrow triangle-shaped areola, and lateral carinae above each spiracle,

rugose around spiracles and inside areola. Abdomen: first tergum as long as

apical width, strigate except medially at base; second tergum strigate,

suturiform atriculation scrobiculate; rest of terga smooth; ovipositor shorter

the length of body, curved down at apex. Legs: fore tibia with 6 spines

along anterior edge, 7 spines at apical edge; fore tarsus l-l/2 times as long

as fore tibia; hind femur 3-1/3 times as long as wide, widest in middle.

Marsh: Megaloproctus (Braconidae) OTF

Wings: recurrent vein distinctly entering first cubital cell at apex; nervulus

postfurcal by 3/4 its length; radiella straight to wing margin.

Maile: Unknown.

Holotype: Female. Turrialba, Costa Rica, June 23, 1948, Franz Shrader.

Deposited in U.S. National Museum, Washington.

This species is easily distinguished by its entirely yellow orange thorax and

abdomen and by the strigate abdominal terga |]-3.

REFERENCES

Brullé, A. 1846. In Lepeletier, Histoire naturelle des insectes: Hyménop-

teres. Paris, Roret vol. 4, 689 pp.

Cameron, P. 1887. Family Braconidae. In Biologia Centrali-Americana.

Insecta Hymenoptera I: 312-419.

Enderlein, G. 1912. Zur Kenntnis der Spathiinen und einiger verwandter

Gruppen. Arch. Natugesch. 78(A)(2): 1-37.

Enderlein, G. 1920. Zur Kanntnis aussereuropaischer Braconiden. Arch.

Naturgesch. 84(A)(ll): 51-224.

Fischer, M. 1980. Zur Taxonomie der Gattungen Hybodoryctes Szépligeti

und Megaloproctus Schulz (Hymenoptera, Braconidae, Doryctinae).

Sitzungsberich. Osterr. Akad. Wissensch. Mathem.-naturw. Kl. Sec. I, Vol.

189, No. 1-3: 1-28.

Harris, R. A. 1979. A glossary of surface sculpturing. Occ. Papers, Calf.

Dept. Food and Agric. No. 28, 31 pp.

Holmgren, A. E. 1869 (1868). In Konglia Svenska Fregatten Eugenies Resa

omkring Jorden. Vetenskapliga Iakttagelser, ii Zoologi; 1, Insects, Haft

12 Hymenoptera, pp. 391-442.

Roman, A. 1910. Notizen zur Schlupfwespensammlung des schwedischen

Reichsmuseums. Ent. Tidskr. 31: 109-196.

Roman, A. 1924. Wissenschaftliche Ergebnisse der schwedischen entomol-

ogischen Reise des Herrn Dr, A. Roman in Amazonas 1914-15, 10.

Hymenoptera: Braconidae, Cyclostomi pro p. Ark. Zool. 16: 1-40.

Schoenherr, C. J. 1838. Gen. et Spec. Curc. 4(2): 868. Roret, Paris.

Schulz, A. A. 1906. Spolia Hymenopterologica. Paderborn, Jungfermann,

356 pp.

Schulz, W. A. 1911 (1909). Zweihundert alte Hymenopteran. Zool. Ann. 4:

1-220. |

Shenefelt, R. D. and P. M. Marsh. 1976. Hymenopterorum Catalogus, Part

13, pp. 1264-1424: Braconidae Part 9, Doryctinae. W. Junk, The Hague.

Szépligeti, G. V. 1902. Tropische Cenocoeliden und Braconiden aus der

Sammlung des Ungarischen National-Museums. Term. Fuz. 25: 39-84.

Szépligeti, G. V. 1904. Sudamerikanische Braconiden. Ann. Mus. Natl.

Hung. 2: 173-197.

315 Contribs Amer. Ent. Inst.,- vol..20, | 1983

Sesser ess

oer,

Figs. 1-8. Wings of Megaloproctus species: 1, castaneus (Br, eras

harpactorinus (End.). 3, platyantennus, n. sp. 4, striatus, n. sp. a

strongylogaster (Cam.). 6, schunckei, n. sp. 7, pertinax (Can.

8, brasiliensis (Szép.).

Marsh: Megaloproctus (Braconidae) 379

é et sse

RESTA ALD

Figs. 9-15. Head, dorsal view, of Megaloproctus species: 9, striatus,

n. sp. 10, harpactorinus (End.). 11, platyantennus, n. sp. 12, castaneus

(Br.). 13, didymus (Br.). 14, brasiliensis (Szep.). 15, schunckei, n. sp.

Figs. 16-19. Propodeum and abdomen of Megaloproctus species: 16,

didymus (Br.). 17, striatus, n. sp. 18, platyantennus, n. sp. 19, brasiliensis.

380 Contrib. Amer. Ent. Inst., vol. 20, 1983

Szépligeti, G. V. 1906. Braconiden aus der Sammlung des Ungarischen

National Museums, I. Ann. Mus. Natl. Hung. 4: 547-618.

Viereck, H. L. 1914. Type species of the genera of ichneumon flies. Bull.

U.S. Natl. Mus. 38: 1-186.

Figs. 20-24. Face of Megaloproctus species: 20, didymus (Br.). 2,

castaneus (Br.). 22, brasiliensis (Szep.). 23, harpactorinus End.).

24, platyantennus, n. sp/

Figs. 25-26. Fore tibia of Megaloproctus species: 25, harpactorinus

(End.). 26, didymus (Br.).

EINE NEUE ART DER GATTUNG HERPESTOMUS WESMAEL, [1845], AUS

ASIEN

(HYMENOPTERA, ICHNEUMONIDAE, PHAEOGENINTI)

Erich Biller

Zoologische Staatssammlung Munchen

Maria-Ward-Strasse 1b, D-8000 Munchen: 19

West Germany

Abstract

A new species of Phaeogenine Ichneumonidae, Herpestomus

henrytownesi, is described from Burma and Pakistan, recording

the genus for the first time from Asia.

Zusammenfassung

Eine neue Art der Subfamilie Ichneuomoninae, der Tribus Phaeo-

genini und der Subribus Phaeogenina wird aus Asien beschrieben.

Fur Pakistan und Burma ist es der erste Nachweis des Vorkommens der

Gattung Herpestomus Wesmael, 1845 .

Einleitung

In der orientalischen Region, in Asien, ist die Tribus Phaeo-

genini (= Ichneumoninae cyclopneusticae, Phaeogeninae, Alomyini

auctt.) bis heute durch verhaltnismagig wenige Arten belegt. Der

Grund daftir dtirfte weniger die mogliche Seltenheit oder versteckte

Lebensweise der Arten und Individuen in diesen Gebieten sein,

vielmehr wurde in den asiatischen Landern, eine Ausnahme bildet

Japan, kaum nach disen Tieren gesucht, zumal auch nur sehr wenige

Entomologen die Gelegenheit hatten dort intensiv Ichenumonidae zu

sammeln.

Es liegen zuer Bearbeitung 3 Exemplare der zu beschreibenden

Art vor. Ein Weibchen und Mannchen wurde in Pakistan gesammelt und

das dritte Exemplar, ein Weibchen, stammt aus Burma. Alle Tierre

enstammen der montanen Region der angefuhrten Landern. Es ist zu

vermuten, daB die Art in den hoheren, bergigen Lagen eine weitere

Verbreitung hat, die angezeigten Lebensraume der disjunten

Verbreitung berechtigen zu diesem SchluB.

Die neue Herpestomus-Art wird als Ausdruck der Hochachtung und

des Dankes fur die fundamentale Arbeit der Enforschung der

Ichneumonidae durch Herrn Professor Doktor Henry Townes aus AnlaB

seines siebzigsten Geburtstages mit seinem Namen belegt.

HERPTESTOMUS HENRYTOWNESI, n. sp. (Abbildungen 1,3)

Weibchen: 6,5-7,5 mm. Kopf glanzend und dicht punktiert, uber

den Antennae dichter querstreifig punktiert; Supraclypealarea

erhaben und vom Clypeus, der ebenfalls stark erhaben ist und apikal

zue einer beinahe geraden Flache stark abfallt, durch eine

deutliche Furche getrennt, zu den Facialorbitae ist die Supra-

clypealarea nur durch die starke Erhabenheit abgesetzt; Occiput

weniger stark punktiert und dadurch glanzender; Malarraum kurzer

381

382 Contrib: Amer. Ent. Inst., vol. 20, 1983

als die Mandibelbasisbreite; der obere Mandibelzahn ist wesentlich

langer als der untere Zahn; Tempora nach hinten verschmalert; die

Genalcarina trifft die Oralcarina kurz vor der Mandibelbasis, die

Oralcarina zwischen Genalcarina und Mandibelbasis kaum erweitert.

Flagellae 32-34 Glieder, das Glied nach dem Postanellus kaum

kurzer als der Postanellus, Flagellae in der Mitte kaum verdickt,

zu den Flagellaeended spitz auslaufend. Thorax glanzend und dicht

punktiert, Pronotum schwach punktiert und grogflachig glanzend, die

Querfurche des Pronotums ist in der Mitte unterbrochen, Praepectus

und Mesopleurum sehr dicht punktiert; Notauli stark eingepragt,

uber ein Drittel des Mesoscutums einnehmend; Sternauli tief

eingepragt und etwa die Halfte des Mesopleurums einnehmend;

Scutellum flach, nur an der Basis gerandet. Nervellus nahezu

oppositus. *Propodeum glanzend und punktiert; Area superomedia

glanzend und unregelmagig langsgerillt, nur in der Anlage penta-

gonal und briet, die Basalcarina jedoch nicht nur eine Ecke bildend,

sondern auch kurz zweickig; Area dentipara quergerunzelt mit schwach

ausgebildeten Apophysen; Area posteromedia schwach ausgehdohlt und

quergestreift. Beine schwach chagriniert und gladnzend; Die

Hintercoxae mit einer + deutlich sichtbaren Leiste die in einen

undeutlichen Tuberkel endet. Abdomen gl&nzend und sehr dicht

punktiert (Abb. 1), ab Abdominaltergit 4 wird die Punktierung

flacher und weniger und die Tergite werden glatt und glanzender;

Postpetiolus breit und dicht punktiert; Gastrocoeli deutlich und

ganz an der Basis des zweiten Tergites liegend, der trennende

Mittelraum breit; der Ovipositor etwas uber das Hinterleibsende

ragend (Abb. 1).

Farbung: Schwarz. Geld sind: Supraclypealarea, Clypeus,

Mandubulae, mit Ausnahme der Zahne, Palpi, Scapi und Pedicelli

ventral, der Oberrand des Pronotums, Subtegulargrat, + das

Scutellum, Flugelbasis, Coxae und Trochantelli der Vorderund

Mittelbeine, die Hintercoxae basal dorsal, die restlichen Hinter-

coxae sind rotbraun. Rotbraun sind die Beine mit Ausnahme der

schwarzen 5. Tarsensegmente und ein + schwarzer Dorsalfleck apikal

auf den Hinterfemora, wie die schwarzen Enddrittel der Hintertibiae

und Hintertarsi. Die Kaudalrander der Tergite konnen getont sein.

Mannchen: 6mm. Es entspricht in der Struktur und Farbung

dem Wiebchen. Flagellae 29 Glieder. Das Gelb des Gesichtes ist

bist zu den Facialorbitae ausgedehnt; die Hintercoxae sind

schwarzlich und basal ohne gelben Dorsalfleck.

Verbreitungsdaten: Holotypus: F, Upper Burma:Mt. Popa, 600-

1000 m, X.— XI. 1937, leg G. Heinrich (Zoologische Staatssammlung

Munchen). Paratypen: 1M, 1 F, West-Pakistan: Kalachitta Range

(bei Basal) °15.7,1966.

Herpestomus henrytownest, n. sp. ist mit Herpestomus brunnt-

ecornts (Gravenhorst, 1829) in verwandtschaftliche Beziehung zu

setzen. Grundsdtzlich unterscheidet sich Herpestomus henrytownest,

n. sp. von Herpestomus brunnicornts (Gravenhorst, 1829) durch

folgende wichtige Punkte: Die Punktierung des Kérpers ist tiefer

und dichter, der Clypeus ist apikal nicht so stark abfallend aus-

gehohlt, die Fuhlerglieder sing langer (Abb. 3,4), der Postpetiolus

ist kraftiger und starker punktiert (Abb. 1,2). Herpestomus

henrytownest, n. sp. hat im Gegensatz zu Herpestomus brunntcornts

(Gravenhorst, 1829) mehrere gelbe Farbmarkierungen.

M= Male. Fo =) Female .

Diller: Herpestomus (Ichneumonidae) 383

LITERATUR

HEINRICH, G. H. 1965. Die Burmesischen Ichneumoninae I.

Entomol. Te. Arg. 86 (8H. -la2ja70sT30.

TOWNES, H., TOWNES, M. & GUPTA, -V.-k%.. 1961. A catalogue and

reclassification of the Indo-Australian Ichenumonidae. Mem.

Amery. Ent. Inst. L252;

4 D Ms 9

2°99 90 29 7 25]9 0/9 5%

I20C¢9

8 oS

SLLLOLEA KAA SI CLL -

Abbildungen 1-4. 1, Herpestomus henrytownesi, F Abdomen.

2, H. brunnicornis F Abdomen. 3, H. henrytownesi, F Kopf

dorsal. 4, H. brunnicornis, F Kopf dorsal.

TOWNESILITUS,A NEW GENUS FOR A SPECIES GROUP IN MICROCTONUS

(HYMENOPTERA: BRACONIDAE, EUPHORINAE) |

E. Haeselbarth

Lehrstuhl fur angewandte Zoologie

der Ludwig-Maximilians-Universitat, Munich

and

Conrad Loan

Ottawa Research Station

Agriculture Canada, Ottawa KIA 0C6

Abstract

Townesilitus, new genus, is described for a small group of Palearctic and Nearctic

species in Microctonus Wesmael characterized by an unusually wide and weakly convex

clypeus, and by the ventral margins of the petiole fused with the sternite.

Muesebeck (1936) reviewed the known Euphorine genera and defined Microctonus

Wesmael by a combination of characters including nongeniculate antennae with a short scape;

two cubital cells, the first confluent with the first discoidal and a complete submediellan

cell; and strongly exserted ovipositor. He pointed out that Microctonus belongs to the same

natural group as Perilitus Nees and he retained the division between these two genera (by

"separation or confluence of the first cubital and first discoidal cells") merely for practical

reasons. During a revision of the European species, the first author found other characters

to divide the Perilitus-Microctonus complex into more natural subgroups. Though the revision

is not advanced enough to decide if Microctonus is a valid genus or only a synonym of

Perilitus, it allows the segregation of a group of species which differs from the others by

morphological and ethological characters. The authors believe these characters are

important enough to justify the erection of a new euphorine genus. In most species of

Microctonus (as usually defined and including the genotype Perilitus aethiops Nees) the

clypeus is narrow and distinctly convex and the sternite of the first abdominal segment is

free from the tergite, i.e. the ventral margins of the petiole are widely separated in front of

the spiracles (Figs. 4 - 6). However, in the above mentioned group, the clypeus is unusually

wide and almost flat, and on the petiole the ventral margins of the tergite are completely

fused with the sternite (Figs. | - 3). The oviposition habits of the species in this group are

characteristic.

We wish to dedicate this new genus to Henry Townes in affectionate admiration for a

great man and an eminent entomologist, and call it Townesilitus, to indicate also the close

relationship of this group to the genus Perilitus Nees.

TOWNESILITUS, new genus

(Pigs. 1-¢)

Front wing 1.5 - 3.0 mm long. Clypeus very broad, at least twice as wide as long, and

only very weakly convex, its apex broadly truncate. Tip of mandible not conspicuously

twisted, bifid, the lower tooth much shorter (ca. | : 4) than the upper one. Occipital carina

complete, dorsally neither weakened nor dipped. Thorax moderately stout, about 2.0 times as

long and 1.5 times as high as broad (measured between the tegulae). Femora, especially

femora | and 2, a little clubshaped (only very slightly, but more so than in most

Microctonus). Sternum I fused with tergum I, reaching about as far back as the spiracles are

located. Petiolus (segment I) strongly arched in side view, its dorsal face longitudinally

striate and laterally carinate from base to apex without a medical convergence of carinae in

front of spiracles and without dorsal pits. Terga Il ff. smooth and shining and with only very

few hairs. Ovipositor exserted, slightly downcurved, but when concealed in its sheaths often

appearing almost straight.

Type-species: Microctonus bicolor Wesmael.

In this genus are included, besides the genotype from Europe: Microctonus deceptor

Wesmael, M. breviradialis Tobias and several undescribed or undetermined species from the

Palearctic region; Microctonus crepidoderae Loan, M. cucumeridis Loan, M. psylliodis Loan

and a small number of undescribed species from the Nearctic region.

Habits: Adult chrysomelid Coleoptera are recorded as hosts in Europe (Sommer 1980)

and North America (Loan 1969). The oviposition behaviour of M. bicolor, psylliodis and

crepidoderae (Loan 1967) is distinctive and similar to the blacine Pygostolus falcatus (Hal.)

oan and Holdaway 196la): the female leaps onto the elytra of the host and, facing the

thorax and parallel to the body of the host, immediately inserts the ovipositor into the apical

384

Haeselbarth & Loan: Townesilitus (Braconidae) 385

region of the haemocoel. While various oviposition positions are know for Microctonus as

here restricted, e.g. the host parasitized through the base of an antenna (Freeman 1967) or

more commonly at the apex of the abdomen (aethiopoides Loan (Loan and Holdaway 1961b),

in all examples the parasite does not mount the host.

Acknowledgements

The authors thank G. Stuart Walley, Winding Way, Nepean, Ontario for his useful

criticisms of the typescript, and Gene Bisdee, Biosystematics Research Institute,

Agriculture Canada, Ottawa for the SEM micrographs,

References

Freeman, B.E. 1967. The biology of the white clover seed weevil Apion dichroum

Bedel (Col. Curculionidae). J. appl. Ecol 4: 535-552.

Loan, C.C. 1967. Studies on the taxonomy and biology of the Euphorinae

(Hymenoptera: Braconidae). IL Host relations of six Microctonus species. Ann. ent.

Soc. Amer. 60: 236-240.

Loan, C.C. 1969. A summary of species of Microctonus north of Mexico with five

new species. Proc. ent. Soc. Wash. 71: 404-416.

Loan, C.C. and F.G. Holdaway. 196la. Pygostolus falcatus (Nees) (Hymenoptera:

Braconidae) a parasite of Sitona species (Coleoptera: Curculionidae). Bull. ent. Res.

D2 473-488.

Loan, C.C. and F.G. Holdaway. 196l1b. Microctonus aethiops Nees (Auct.) and

Perilitus rutilus Nees (Hymenoptera: Braconidae) European parasites of Sitona weevils

(Coleoptera: Curculionidae). Can. Ent. 93: 1057-1059.

Muesebeck, C.F.W. 1936. The genera of parasitic wasps of the braconid subfamily

Euphorinae with a review of Nearctic species. USDA Misc. Bull 241.

Sommer, G 1981. Biologie and Parasitenkomplex der Halticen Gattung

Phyllotreta, Ph.D. Thesis, Albert-Ludwigs University of Freiburg.

386 Cent rib. Amer. Ent. Insti vol. 20, 1983

Figs. 1-3. SEM micrographs of Townesilitus bicolor: 1-2, Dorsal and

ventral views of tergite 1 respectively. 3, Clypeus.

Haeselbarth & Loan: Townesilitus (Braconidae) 387

Figs. 4-6. SEM micrographs of Microctonus vittatae: 4-5, Dorsal

and ventral views of tergite 1 respectively. 6, Clypeus.

HOST AND GENERIC RELATIONS OF THE EUPHORINI

(HYMENOPTERA: BRACONIDAE)

Conrad Loan

Ottawa Research Station, Agriculture Canada, Ottawa K1A 0C6

Abstract

The tribe Euphorini as presently constituted has 2 major groups of genera, the parasites

of nymphal Heteroptera and Psocoptera and the parasites of adult insects. The tribe is here

restricted to those genera parasitic on nymphal hosts and a new tribe, Perilitini (type genus

Perilitus Nees 1818) is proposed for the parasites of adult insects. Keys are given to the

Nearctic genera of each tribe. The relationship of structural modifications of the ovipositor

and petiole with host specificity and method of egg deposition is discussed. The genera of

both tribes have simple life cycles closely integrated with their hosts and synchronized with

the seasonal occurrence of the life stage attacked.

The Nearctic Euphorini as presently constituted (Marsh in Krombein et al 1979) comprise

15 morphologically heterogeneous, small genera with the greatest number of species in

Microctonus Wesmael and Peristenus Foerster. The group attacks a wide range of hosts in

adult insects, and nymphs of various Heteroptera and Psocoptera. In this paper, the generic

and host relations are discussed and a new tribe is proposed for the parasites of adult

insects. Since Muesebeck (1936) revised the subfamily Euphorinae, Mason (1964) reported

Chrysopophthorus Goidanich, and Shenefelt (1960), Ropalophorus Haliday for the Nearctic

region. The Nearctic genus Holdawayella Loan, and the Holarctic Townesilitus Haeselbarth

and Loan were described (Loan 1967, Haeselbarth and Loan 1983). The name Ropalophorus

Haliday was validated for Eustalocerus Foerster (Hincks 1943), Leiophron Nees for Euphorus

Nees (Muesebeck 1958) and for Euphoriana Gahan (Loan 1974), and Peristenus Foerster for a

group of species removed from Leiophron (Loan and Bilewicz-Pawinska 1973).

The parasites of Heteroptera and Psocoptera show considerable diversity in certain

structural features; however, the genera they represent appear to form anatural group. This

is indicated by Muesebeck (1936) who noted that the ovipositor of nymphal parasites

(Euphorus, Euphoriana, Aridelus) was subexserted but that it was prominently exserted in

parasites of Coleoptera. The structure of the ovipositor, and possibly also of the petiole,

indicates the method of egg deposition: if subexserted and strongly decurved, the host is

grasped and held for oviposition, and if exserted, whether in the Euphorini or Perilitini, the

host is parasitized by the parasite striking at it from various positions without grasping it.

Peristenus and Leiophron, characterized by a subexserted, strongly decurved ovipositor and a

short petiole chase the host nymph, and grasp and lift it from the substrate and insert the egg

into the ventral area of the abdomen (Brindley 1936, Glen 1972). In this example, both the

conformation of the ovipositor and of the petiole fit the requirements for oviposition in a

small, active host. Little is known of Aridelus, but the unusually long and slender petiole

may have developed to permit the gaster to reach around and under the dorsoventrally

compressed abdomen of the nymph for oviposition. The exserted apical part of the

ovipositors of the other parasites of Heteroptera is very short, and straight, and I have seen

both Holdawayella and Wesmaelia strike repeatedly at a host nymph, often without success

as no parasite eggs were recovered from the host. The tingid host of Holdawayella is

gregarious and sedentary requiring little movement by the parasite to locate and parasitize

it. The host is also spinose, and to oviposit the female parasite, in a lethargic manner,

partly mounts the nymph and bends the gaster under its thorax or abdomen. On the other

hand, the Nabis host of Wesmaelia is active and searches for prey, and it is parasitized by the

parasite facing and striking at the side of the nymphs's abdomen, a method similar to that of

Microctonus. The long petiole of Wesmaelia may allow it to oviposit at a maximum distance

from its predaceous host.

Euphorines with prominent ovipositors attack adult hosts and form a second natural

group of genera: 6 of the 9 genera are parasites of Coleoptera. These 6 genera are less

diverse than the parasites of nymphs though the flagellum of Ropalophorus and Streblocera is

highly specialized and the wing veins of Cryptoxilos are greatly reduced. Microctonus is the

best known genus and it is clearly closely allied with Perilitus in structure and biology. The

oviposition behavior of Microctonus has the parasite facing the host with the gaster bent

downwards and forwards between its legs and the ovipositor extruded past its face; the

ovipositor is then thrust into a specific area of the host's body. The ovipositor of Microctonus

is straight or weakly decurved whereas that of the closely allied genus Townesilitus is

distinctly decurved, which is more efficient because when the female leaps onto the back of

a host to oviposit, the gaster is bent downwards and the ovipositor is inserted in the apical

area of the abdomen. The petiole of Townesilitus is the only example among parasites of

adult Coleoptera which has the ventral margins of tergite | fused with the sternite.

388

Loan: Euphorini (Braconidae) 389

This structural modification may have developed to give greater strength to the gaster during

the downward thrust of the ovipositor than it would have if the ventral margins of tergite 1

were separated from the sternite.

Much of the heterogeneity in the present tribe Euphorini results from a grouping of

nymphal parasites with those of adult insects. This anomaly would be resolved, at least in

part, by recognition of these apparently natural groups as representing different tribes.

Accordingly, it is here proposed to erect a new tribe, the Perilitini (from the oldest genus

Perilitus Nees 1818), to include those genera whose parasites attack adult insects and to

restrict the parasites of nymphs to the tribe Euphorini. The Nearctic tribes of the

Euphorinae and the genera of Euphorini and Perilitini are determined in the following keys.

KEY TO NEARCTIC TRIBES OF THE EUPHORINAE

1. Tergite 1 sessile or subsessile with spiracles near base. Parasites of adult Scolytidae

ssdanpasiasaneonene: STOO NOTIN

Tergite 1 petiolate or subpetiolate or long and slender with spiracles at or slightly

DEHING ATLAS ..ecosscnanavepesonie Ze

2. Tergite 1 with ventral margins free from sternite, strongly petiolate. Lateral margins of

tergite 2 carinate. Second cubital cell closed and medius present and straight. Cocoon

pensile. Parasites chiefly of larvae of Lepidoptera ...........cee.000. Meteorini

Tergite 1 with ventral margins free from or fused with sternite, subpetiolate or long and

slender. Lateral margins of tergite 2 non-carinate. Second cubital cell, if present, open

and medius present and bowed, or second cubital cell closed and medius affaced.

Cocoon not. pensile, in: Sil 's ssesecssadsntsasees Os

3. Ovipositor subexserted or very short (sometimes extruded in death) with sheaths not

longer than scape combined with pedicel. Petiole short or if unusually long then as long

as gaster behind petiole. Front wing with strong medius and 3 cubital cells, or weak or

effaced medius and |- 2 cubital cells. First abscissa of cubitus usually present but if

absent then ventral margins of tergite | not fused with sternite or gaster not strongly

laterally compressed. Parasites of nymphs of Heteroptera and Psocoptera

Khesintthinhieares CUDOOTIN

Ovipositor prominently exserted with sheaths greatly longer than scape combined with

pedicel. Petiole usually short, but if unusually long and slender then not as long as

gaster behind petiole. Front wing with weak or effaced medius and 2-3 cubital cells and

first abscissa of cubitus present or absent, or strong medius and first abscissa of cubitus

present or absent. Parasites of adult Coleoptera, Hymenoptera and Neuroptera or host

UNKNOWN) sscascavcsesaveseese HOM tL

KEY TO NEARCTIC GENERA OF THE TRIBE EUPHORINI

1. Front wing with 3 cubital cells (Fig. 5). Parasites of Pentatomidae

eeccccccceseseceeeee Aridelus Marshall

Front wing with 1 or, more usually, 2 cubital Cells ..sscsccsosesesneees Ze

2. Tergite 1 with ventral margins widely separated from base to apical margin, or ventral

margins meeting only at base. Ovipositor subexserted and strongly decurved

Tergite 1 with ventral margins fused with sternite, or apparently fused but with margins

nearly touching or one margin overlapping the other. Ovipositor subexserted or very

short and straight or weakly AeCUrVed c.ccccccccccccccceee 4.

3. Ventral margins of tergite | widely separated from base to apex. Parameres of male

genitalia extend basally on ventral side and longer than aedeagus. Front wing veins

usually reduced and nervellus often incomplete or absent in hind wing (Fig. 6). Parasites

Of Miridae ....cccccccsecceeeee Leiophron Nees

Ventral margins of tergite | usually meeting at base and opening posteriorly. Parameres

of male genitalia about as long on dorsal side as on ventral and not as long posteriorly as

aedeagus. Apart from effaced medius, wing veins not reduced and nervellus always

present (Fig. 7). Parasites of Miridae ...........0.00... Peristenus Foerster

4, Front wing with 2 cubital cells and complete radius. Nervellus present. Petiole long and

slender, thickened medially. Ovipositor straight (Fig. 8). Parasites of Nabidae

e cvccccecccosseccoee WESMAElia Foerster

Front wing with | or 2 cubital cells and incomplete or effaced radius. Nervellus absent

or weakly indicated at base. Petiole slender but not unusually long and not thickened

medially. Ovipositor straight or weakly decurved. Smaller parasites ........ccccccecsees De

5. Ventral margins of tergite 1 apparently fused but margins slightly apart or one margin

overlaps the other. Wing veins greatly reduced with only basal, submedius, cubitus,

nurvulus basally present in front wing. Ovipositor barely perceptible, weakly decurved,

(Fig. 9). Parasites of Psocidae .......ccccccceseeee Euphoriella Ashmead

Ventral margins of tergite 1 fused with sternite. Wing veins less reduced but second

390 Contrib. Amer. Ent. Inst., vol. 20, 1983

abscissa of radius incomplete approaching wing margin. Ovipositor straight (Fig. 10).

Parasites Of Tingidae ........c.cccccccseee Holdawayella Loan

KEY TO NEARCTIC GENERA OF THE TRIBE PERILITINI

1. Medius weak basally or effaced. Parasites of Hymenoptera, Neuroptera or host unknown

Medius strong. Parasites of Coleoptera .scccccsccsocccceeee 4e

2. Front wing with 3 cubital cells. First abscissa of cubitus and nervellus present. Petiole

cylindrical, unusually long and slender, about as long as thorax and propodeum (Fig. by).

Parasites Of Chrysopidae .....ccccsccesseeeee Chrysopophthorus Goidanich

Front wing with 2 cubital cells. First abscissa of cubitus absent, nervellus present or

absent. Petiole cylindrical or dorsoventrally compressed, shorter than thorax and

PFOPOCEUM .« cccoccccccccccceecs de

3. Gaster strongly laterally compressed. Eyes of female very large and prominent. Petiole

dorsoventrally compressed, ventral margins of tergite | not fused with sternite, dorsal

face with deep and long glymmae. Nervellus present (Fig. 12). Host unknown

ssececscscesecseeeees Myiocephalus Marshall

Gaster not unusually laterally compressed. Eyes not unusually prominent. Petiole

cylindrical with ventral margins of tergite | fused with sternite, dorsal face without

glymmae. Nervellus absent (Fig. 13). Parasites of Bombidae, Ichneumonidae

sesedesetveseseoscas ONTItre TUS Marshall

4, Flagellum short with 8- 12 articles, unmodified or proximal and distal articles elongated

ANd SCAaPe SHOIt ...cccccccccccssesee De

Flagellum not unusually short with at least 15 articles, unmodified or geniculate with

basal articles specialized or simple. Scape elongated Or SHOrt .rccccccccscsceseees 6.

5. Flagellum clavate with proximal and distal articles elongated. Wing veins relatively

complete (Fig. 14). Parasites of Scolytidae .......sceseeeeeeeee Ropalophorus Haliday

Flagellum non- clavate. Wing veins reduced with recurrent, subcostella, submediella,

and nervellus absent (Fig. 15). Parasites of Scolytidae .....cccsessssseseee Cryptoxilos Viereck

6. Flagellum geniculate. Scape elongated and basal flagellar articles specialized by

structure and pubescence, or non- specialized. Wing veins complete except first

abscissa of cubitus absent (Fig. 16). Parasites of Chrysomelidae

secsccscscssesseeses Streblocera Westwood

Flagellum nongeniculate. Scape short and basal flagellar articles unmodified

7. First abscissa of cubitus present (Fig. 17). Parasites of Coccinellidae, Curculionidae,

Morabinae (Orthoptera) .....c..cccccccceoeee Perilitus Nees

First abscissa of CUbItUS ADSENT .recccccccseseececes Se

8. Clypeus longer than wide and strongly convex. Tergite | with ventral margins widely

separated between base and apical margin. Female oviposits from the substrate without

leaping onto back of host (Fig. 18). Parasites chiefly of Chrysomelidae and

Curculionidae ......ccccccccsceeee Microctonus Wesmael

Clypeus unusually wide, at least 2.0 times as wide as long and almost flat. Petiole

cylindrical in front of spiracles with ventral margins of tergite | fused with sternite.

Female leaps onto back of host to oviposit (Fig. 19). Parasites of Chrysomelidae

ssescsscesssssseeeee LOWNESilitus Haeselbarth and Loan

Muesebeck (1963) briefly outlined the host relations of the Nearctic genera of

euphorines, demonstrating that each genus attacks a specific host group. Perilitus Nees,

however, is a unique exception as it not only attacks weevils and leaf beetles in Europe

(Jackson 1928, Loan and Holdaway 1961, Waloff 1961) but it is also a parasite of morabine

grasshoppers in Australia (Blakith 1967). Hosts are known for 10 of the Nearctic genera and.

inferred for 3 others from Palearctic records (Aridelus Marshall, Capek and

Davidova-Vilimova 197 8; Perilitus loc cit; Syntretus Foerster, Cole 1959, Alford 1968) and |

from the Orient (Streblocera Westwood, Watananbe 1942). The host of Myiocephalus Marshall

is unknown. The associations are listed in Table 1 with the stage attacked and the stage

killed. The tribe Euphorini, in the restricted sense proposed in this paper, has almost as

many genera and nearly twice as many species as the tribe Perilitini.

The known parasite life cycles are closely integrated with their host life histories and

modifications are found more in the genera attacking nymphs than in the parasites of

Coleoptera. Parasites (Euphoriella, Leiophron, Peristenus) whose hosts (psocids, plant bugs)

overwinter as eggs pass the winter as diapausing adults in larval cocoons and emerge to

attack early instar nymphs (Fig. |) (psocid parasites Sommerman 1956; plant bug parasites

Waloff 1967, Loan 1980). The parasite cocoons overwinter in soil, but Ihave also found

Euphoriella cocoons beneath tree bark. Other parasites overwinter as first instar larvae in

adult hosts though Chrysopophthorus is exceptional because it overwinters in the larval

Loan: Euphorini (Braconidae) 301

cocoon, stage unknown (Raymond Manuel in litt). The tingid parasite Holdawayella attacks

fifth instar nymphs that are mature by mid-summer and synchronizes with them by slow

development of the overwintered larva and of the cocoon stages (Fig. 2) (Loan, Gerber and

Reid, 1971). Microctonus colesi Drea, a parasite of the alfalfa weevil Hypera postica Gyll,

shows a remarkably similar adaptation to its host. This, the only known species of

Microctonus to exclusively parasitize larvae attacks late instar hosts and the first instar

parasite larva diapauses in the adult host and over winters; thus its life cycle corresponds

closely with that of Holdawayella (Fig. 2). Wesmaelia has 2 broods of parasites in the season

which attack fifth instar nymphs in early and late summer (Fig. 3). At Ottawa, I have reared

larvae of Wesmaelia as early as April, and recovered eggs laid by the second brood of

parasites as late as September. The genus Aridelus has only been reared in the Palearctic

region. Capek and Davidova-Vilimova (1978) identified larvae of A. egregius

(Schmiedeknecht) from early and late instar nymphs of the pentatomid Captosoma. They

suggest that the first instar larva overwinters; the oviposition behavior of A. egregius was

not obser ved.

Life-histories of 3 of the genera attacking adult Coleoptera, Microctonus, Perilitus and

Townesilitus are well known (Microctonus, Nearctic records in Loan 1969; Perilitus, Balduf

1926 Townesilitus, as Microctonus psylliodis Loan, M. cucumeridis Loan, M. crepidoderae

Loan, loc cit). The life cycles of these parasites (Fig. 4) and that of the Nabis parasite

Wesmaelia closely agree with the chief difference that the first brood of parasites of the

former attack overwintered, sexually active hosts. Of the euphorines parasitic on other adult

insects, Syntretus has not as yet been reared from a Nearctic host but, in Europe, species are

recorded from ichneumonids (Cole 1959) and bumble- bees (Alford 1968). In New Jersey in

September 1970, numerous Syntretus, presumably searching for hosts, were captured in black

light traps (James Stewart in litt). Thus the species may overwinter in adult hosts as first

instar larvae. The life cycle of Chrysopophthorus appears similar to that of Microctonus

with 2 broods of the parasite attacking the bivoltine host (Raymond Manuel in litt).

Acknowledgements

The modifications to the tribal classification of the Euphorinae developed from

suggestions by W. R. M. Mason, Biosystematics Research Institute, Agriculture Canada,

Ottawa. To him and G. Stuart Walley, Winding Way, Nepean, Ontario, I express sincere

thanks for help with the manuscript. I also wish to thank Raymond Manuel, Department of

Entomology, Macdonald College of McGill University, Montreal for specimens of

Chrysopophthorus and information on its life cycle.

List of figures

1-4, Life cycle outlines. 1, Euphoriella, Leiophron, Peristenus: 2, Holdawayella; 3,

Wesmaelia; 4, Microctonus, Perilitus, Townesilitus.

5-19. Wings. 5, Aridelus; 6, Leiophron; 7, Peristenus; 8, Wesmaelia; 9, Euphoriella; 10,

Holdawayella; 11, Chrysopophthorus; 12, Myiocephalus; 13, Syntretus; 14,

Ropalophorus; 15, Cryptoxilos; 16, Streblocera; 17, Perilitus; 18, Microctonus; 19,

Townesilitus. c igen cate a

Note: Figures 5 - 19 are not to scale and are diagrammatic in area and outline.

References

Alford, D.V. 1968. The biology and immature stages of Syntretus splendidus (Marshall)

(Hymenoptera: Braconidae, Euphorinae) a parasite of adult bumblebees. Trans. R. ent.

Soc. Lond. 120: 375-393.

Balduf, W. V. 1926. The bionomics of Dinocampus coccinellae Schrank. Ann. ent. Soc. Amer.

19: 465-498.

Blakith, R. E. 1967. A hymenopterous primary parasite of morabine grasshoppers. Aust. J.

Zool. 15: 93-102. oe

51-56.

Capek, M., and J. Davidova-Vilimova. 1978. On the life history of Aridelus egregius

(Hymenoptera: Braconidae) a parasite of European Captosoma (Heteroptera:

Plataspidae). Actaentomol. bohemoslov. 75: 243-254.

Cole, L.R. 1959. On anew species of Syntretus Foerster (Hym., Braconidae) a parasitie on

an adult Ichneumonid, with a description of the larva and notes on its life history and

that of its host, Phaeogenes invisor (Thunberg). Ent. Mon. Mag. 95: 18-21.

Glen, D. M. 1977. Ecology of the parasites of a predatory bug, Blepharidopterus angulatus

(Fall.). Ecol. Entomol. 2: 47-55.

392 Contrib. Amero Ent. Inst. vol. 20; 1983

Haeselbarth, E., and C. Loan. 1983. Townesilitus, a new genus for a species group in

Microctonus. Contrib. Amer. Ent. Inst. 20 (1): 384-387.

Hincks, W. D. 1943. Nomenclature notes on Braconidae and Aphidiidae (Hym.). Entomologist

76: 101.

Krombein, K. V., P. D. Hurd, Jr., and D. R. Smith. 1979. Catalog of Hymenoptera in

America north of Mexico. Smithsonian Institution Press.

Loan, C. C. 1967. Studies on the taxonomy and biology of the Euphorinae (Hymenoptera:

Braconidae). III. Anew Canadian genus and species parasitic on Tingidae

species (Hymenoptera: Braconidae, Euphorinae). Proc. ent. Soc. Wash. 71: 404-416.

Loan, C. C. 1974. The European species of Leiophron Nees and Peristenus Foerster

(Hymenoptera: Braconidae, Euphorinae). Trans. R. ent. Soc. Lond. 126: 207-238.

Loan, C. C. 1980. Plant bug hosts (Heteroptera: Miridae) of some euphorine parasites

(Hymenoptera: Braconidae) near Belleville, Ontario, Canada. Naturaliste can. 107:

87-93.

Loan, C. C., G. H. Gerber, and D. G. Reid. 1971. Biosystematics of the tingid parasite

Ho awayells in Ontario (Hymenoptera: Braconidae, Euphorinae). Can. Ent. 103:

Loan, C. C., and T. Bilewicz-Pawinska. 1973. Systematics and biology of four Polish species

of Peristenus Foerster (Hymenoptera: Braconidae, Euphorinae). Environ. Ent. 2:

271-278.

Mason, W.R.M., 1964. The genus Chrysopophthorus Goidanich (Hy menoptera:

Braconidae). Can. Ent. 96: 1005-1017. |

Muesebeck, C. F. W. 1936. The genera of parasitic wasps of the braconid sub-family

Euphorinae, with a review of the Nearctic species. USDA Misc. Bull. 241.

Muesebeck, C. F. W. 1958. New Neotropical wasps of the family Braconidae (Hymenoptera)

in the U. S. National Museum. Proc. U. S. Nat. Mus. 107: 405-461.

Muesebeck, C. F. W. 1963. Host relationships of the Euphorini (Hymenoptera: Braconidae).

Proc. ent. Soc. Wash. 65: 306.

ent. Soc. Wash. 58: 149-152.

Waloff, N. 1961. Observations on the biology of Perilitus dubius (Wesmael) (Hymenoptera:

Braconidae), a parasite of the chrysomelid beetle Phytodecta olivacea (Foerster). Proc.

R. ent. Soc. Lond. 36 96-102. ‘

—— = eee 0 eee

Euphorinae) parasitic on Miridae on broom. Trans. R. ent. Soc. Lond. 119: 187-213.

Watanabe, C. 1942. A supplementary note on the genus Streblocera (Hymenoptera:

Braconidae). Ins. Mats. 16: 158.

Loan: Euphorini (Braconidae)

393

Table 1. Hosts of Nearctic euphorine and perilitine genera with stage attacked and stage

killed

Parasite Number Family

Genus Nearctic of

Species Host

EUPHORINI

Parasites of Heteroptera and Psocoptera

Aridelus 3 Pentatomidae

Leiophron 9 Miridae

Peristenus 25 Miridae

Wesmaelia | Nabidae

Euphoriella 10 Psocidae

Holdawayella 2 Tingidae

PERILITINI

Parasites of Hymenoptera and Neuroptera

Chrysopophthorus I Chry sopidae

Syntretus 3 Ichneumonidae

Bombidae

Parasites - host- unknown

Myiocephalus l

Parasites of Coleoptera

Ropalophorus l Scolytidae

Streblocera l Chrysomelidae

Cry ptoxilos | Scolytidae

Perilitus 2 Coccinellidae

Curculionidae

Townesilitus 3 Chrysomelidae

Carabidae

Chrysomelidae

Microctonus 25 Curculionidae

Nitidulidae

Tenebrionidae

Stage of host

Attacked Killed

Early nymph

Early nymp

Early nymph

Late nymph

Early nymph

Late nymph

Adult

Adult/larva

Adult

Nymph and adult

“Nymph and adult

Adult

Nymph and adult

Adult

394 Contrib, Aner. Ent. oinst) vole, 19383

HETEROPTERA MIRIDAE Egg stage overwinters with

PSOCOPTERA PSOCIDAE _ 1 brood of nymphs in life cycle

Parasite adults

emerge April — May

Parasite adults

diapause and over-

‘ ; Attack first and

winter in cocoon

second instar nymphs

EUPHORIELLA

LEIOPHRON

\ PERISTENUS

Cocoons formed in soil

(mirid parasites) or in Larvae emerge from

soil or under tree bark - mature nymphs or

(psocid parasites) Bena adults May — July :

HETEROPTERA TINGIDAE Adult stage overwinters with

1 brood of nymphs in life cycle

Parasite adults

wr emerge July }

Larvae emerge from adult

hosts in June, cocoon HOLDAWAYELLA Attack fifth instar

in soil tingid nymphs

iS First instar larvae

diapause in adult

tingids and over- 9

winter

Figs. 1-2. Life cycle outlines: 1, Euphoriella, Leiophron, Peristenus.

2, Holdawayella.

Loan: Euphorini (Braconidae) 395

HETEROPTERA NABIDAE Adult stage overwinters with

2 broods of nymphs in life cycle

First brood of parasite

adults emerge in May

rag

Parasite larvae develop

in April — May, emerge

and cocoon

Attack nabid nymphs.

First instar larva

diapauses in adult host

and overwinters

WESMAELIA

Second brood of parasites

emerge in July — August

COLEOPTERA CARABIDAE

COCCINELLIDAE

First brood of parasite

adults emerge in May

Parasite larvae develop

in April — May, emerge

and cocoon

Attack sexually-immature

adults of summer generation.

First instar larvae diapause

and overwinter in adult host

MICROCTONUS

PERILITUS

TOWNESILITUS

Te Second brood of parasites

CURCULIONIDAE

CHRYSOMELIDAE NITIDULIDAE

TENEBRIONIDAE

Attack fifth instar

nabid nymphs in May — June

}

Egg and first instar larva

carried into new adult and

develop. Final instar larva

emerges, cocoons in soil

Adult stage overwinters

with 2 broods of adults

in life cycle

Attack overwintered

sexual adults

)

Parasite larva develops,

emerges, cocoons in soil

emerge in June — July

Figs. 3-4. Life cycle outlines: 3, Wesmaelia.

Microctonus, Perilitus, Townesilitus.

396 Contrib. Amer. Ent. Inst., vol. 20, 1983

ARIDELUS FISHERI (VIERECK)

EUPHORINI

LEIOPHRON TRIGONOTYLIDIS LOAN

PERISTENUS PALLIPES (CURTIS)

WESMAELIA PENDULA FOERSTER

HOLDAWAYELLA TINGIPHAGA LOAN

EUPHORIELLA SOMMERMANAE MUESEBECK

Figs. 5-10. Wings: 5, Aridelus. 6, Leipphron. 7, Peristenus. 8,

Wesmaelia. 9, Euphoriella. 10, Holdawayella.

Loan: Euphorini (Braconidae) 397

CHRYSOPOPHTHORUS AMERICANUS MASON

SYNTRETUS VIGILAX (PROVANCHER) eg E R : L L T : Ni :

ROPALOPHORUS WISCONSINENSIS SHENEFELT

GRYPTOXILOS CONVERGENS MUESEBECK

PERILITUS RUTILUS (NEES)

MICROCTONUS VITTATAE MUESEBECK

TOWNESILITUS PSYLLIODIS (LOAN)

Figs. 11-19. Wings: 11, Chrysopophthorus. 12, Myiocephalus. 13,

Syntretus. 14, Ropalophorus. 15, Cryptoxilos. 16, Streblocera. 17, Peri-

litus. 18, Microctonus. 19, Townesilitus.

ICHNEUMOFAUNA DO SUDESTE E SUL DO BRASIL. V. NOVA ESPECIE

DE PHILODRYMUS (ANOMALONINAE, HYMENOPTERA)*

Vineita Grat**

Depto. de Zoologica

Universidade Federal do Parana

Curitiba, Parana, Brasil

Abstract

Philodrymus townesi , n. sp., parasitoid from Hypsipyla grandella (Zeller,

1848) (Phycitinae, Pyralidae), borer of the fruits of Cedrela fissilis Meliaceae,

is described from Manancias da Serra, Piraquara, Parana, Brazil.

Nas chaves e descrigoes de Anomaloninae de Townes (1971: 123-159)

e de Gauld (1976: 22- 29; 81-82( este novo ichneumonideo é um Philodrymus

com algumas diferencas 4 nivel genérico. As pupas de Hypsipyla grandella

(Zeller, 1848), Phycitinae, Pyralidae foram obtidas de frutos do cedro Cedrela

fissilis, Meliaceae) coletados no municipio de Piraquara (Mananciais da Serra),

Paranda, Brasil. Estas quando parasitadas s& parcialmente consumidas pelo

ichneumonideo que faz para sair uma abertura circular na extremidade cefalica

da exivia da pupa. As medidas entre parenteses sao em centésimos de mili-

metro e os indices sa os de Gauld (1976: 8-9). O nome especifico é uma home-

nagem ao prof. Dr. Henry K. Townes.

PHILODRYMUS TOWNESI, N. SP.

FEMEA. Cabega amarela exceto o vértice, a fronte até os alvéolos antenais,

o pés-occipicio até a fossa proboscidial, as extremidades distais das mandibulas

e as antenas, pretas. Tdorax ferrugineo-avermelhado exceto os lobos mediano e

laterais do mesoscuto, a area deprimida do escutelo, a maior parte da mesopleura

e o mesosterno (menos o prepecto e as estreitas areas anterior, posterior e su-

perior da mesopleura), pretos; a propleura amarela. Asas hialinas, lavadas de

marrom-péalido nas partes distais, as veias e os estigmas pretos. Pernas ante-

riores amarelas exceto Os fémures e parte basal das tibias amarelo-ferrugineos

e os tres tarsOmeros distais e as garras enegrecidas; pernas médias amarelas

exceto os trocantelos, os fémures e Area basal das tibias ferrugineos e os tarsos

pretos. Pernas posteriores pretas exceto os esporoées tibiais ferrugineos. Ab-

domen preto exceto os esternitos o partes mais laterais dos tergitos e ovipositor

ferrugineo-amarelados.

Olhos com pubescéncia muito curta.e esparsa, as margens inferiores forte-

mente convergentes, carena occipital completa, médio-dorsalmente convexa e

proxima aos ocelos posteriores, no extremo inferior (carena genal) curva em

arco e unida a base da mandibula; parte inferior da gena muito larga, quase o

dobro da largura da base da mandibula. Fronte com curta carena mediana entre

os alvéolos antenais e acima destes com rugas transversais. Antenas longas mas

mais curtas que 0 corpo, o escapo na extremidade distal fracamente obliquo,

1.5 vezes o comprimento do pedicelo (36:24), o quarto flagelomero com o com-

primento 2.8 vézes a largura. Clipeo liso com pontos esparsos, com un dente

apical mediano; face lisa com pontos menores e mais densos que do clipeo exceto

na. area mediana, sem pontuac¢ao. Mandibulas bidentadas com o 4pice recurvado,

*Contribuigao n° 480 do Depto. de Zoologia da Univeridade Federal do

Parana, Brasil.

**Bolsista do CNPq.

398

Graf: Ichneumofauna 399

o dente inferior bem mais curto que o superior. Pronoto subvertical com sulco

transversal interrompido estreitamente no meio e area superior lisa e brilhante,

margem posterior inferior do pronoto pontiaguda. Mesoscuto abruptamente arre-

dondado anteriormente, com fraca concavidade subapical. Lobos do mesoscuto

com pontuagao pouco densa e intervalos mate. Notaulices deprimidos e com a

regiao médio-posterior do mesoscuto irregularmente rugosos; sutura transversal

do mesoscuto presente. Esternaulo deprimido até a metade da mesopleura com

a parte posterior muito dilatada em alvéolo. Mesosterno e area abaixo do ester-

naulo liso e brilhante com pontuacad fina e pouco densa. Carena pdés-pectal

completa. Tibia média com dois espordes, o externo bem mais curto que o in-

terno (24:30). Troc&uter posterior 2.0 vézes o comprimento do trocantelo, tarso

posterior dos machos nao dilatados; garras tarsais pectinadas até o apice sem

dimorfismo sexual. Asa posterior com 3-4 hamulos. Projec&0 posterior do pro-

pddeo esculpida, com 0.56 do comprimento da coxa posterior, em vista ventral.

Terceiro tergito abdominal com sulco lateral até e espiraculo. Valva do oviposi-

tor 2.3 vezes o comprimento do tergito III; apice do ovipositor afilado com quatro

diminutas carenas distais, o comprimento da parte afilada 0.31 (10:32) da largura

média em vista lateral, a extremidade fracamente curvada para baixo.

Comprimento da asa anterior 9.33 mm e comprimento total aproximado 16.17

mm,

Indices: braquial 2.35 (98:38); disco-braquial 1.21 (160:132); marginal

2.87 (272:92); radial 1.37 (42:32); intercubital 0.55 (16:34); peciolar 7.33

(252:48) e dorso-abdominal 2.19 (324:148).

MACHO: Nono esternito abdominal posteriormente arredondado. Parameros

do edeago esclerotizados com extremidades proximais angulosas, as distais ndo

aleancam o apice do edeago e curvos.

Comprimento da asa anterior 8.25 mm e comprimento total aproximado:

15.25 mm. 7

Indices: braquial 2.12 (68:32); disco-braquial 1.16 (130:112); marginal

2.81 (236: 84); radial 1.36 (38:28); intercubital 0.50 (16:32); peciolar 6.44

(232: 36) e dorso-abdominal 2.14 (300:140).

TIPOS: holdétipo femea e alétipo, BRASIL, Parana, Municipio de Piraquara

(Mananciais da Serra), obtidos em laboratério de pupa de Hypsipyla grandella

(Zeller, 1848), Pyralidae em frutos de cedro (Cedrela fissilis, Meliaceae), em

17, VIII.1976, V. Graf col., na colegéo do Depto. de: Zoologica da Universidade

Federal do Parana.

MATERIAL ESTUDADO: 1 fémea, 3 IX 1976 e 4 machos de 27.VII., 1.1X,

2.IX.e 17, 1X.1976, todos obtidos em laboratério, parasitas de Hypsipyla gran-

della em frutos de cedro, coletados no municipio de Piraquara (Mananciais da

Serra), Paranda, na coleca do Depto. de Zoologica da Universidade Federal do

Povana, 9

DISCUSSAO TAXONOMICA. Em Philodrymus sao reconhecidas as espécies

P. vitticollis (Cresson, 1874) do México, P. minor (Szépligeti;. 1905) do Brasil,

Sao Paulo e P. minor (Szépligeti, 1906) da Bolivia e Peri: (Townes, 1971: 154

e Gauld, 1976:82); diferem da nova espécie na coloragao escura da extremidade

distal da asa anterior em P. vitticollis e P. major e no “comprimento da célula.

discoidal mais longa que a braquial" e a "coloragao do térax amarela com man-

chas" em P,minor. Em relagéo a diagnose de Philodrymus de Gauld (1976:81-82)

a nova espécie difere no seguinte: 1) quarto segmento flagelar com comprimento

acentuadamente menor que o estabelecido para Philodrymus e Podogaster; 2)

carena genal forma um arco na sua parte inferior, préximo a carena hipostomal,

e se une a base da mandibula; as genas sao bem mais largas que as bases das

mandibulas que se prolongam para baixo na parte posterior; na diagnose de

Gauld a carena genal pode estar ausente ou se presente unida a carena hiposto-

mal acima da base da mandibula; 3) pronoto com sulco transversal distinto sé

interrompido estreitamente no meio; na diagnose de Philodrymus este sulco esté

400 Contrib. Amer. Ent. Inst., vol. 20, 1983

ausente; 4) margem inferior posterior do pronoto aguda, enquanto que na diag-

nose, truncada; 5) trocanter das pernas posteriores cerca de 2.0 vezes o com-

-primento do trocantelo, na diagnose 2.5 vezes; 6) valvula do ovipositor (val-

vula 3) com mais que o dobro do comprimento do terceiro tergito, na diagnose

somente 1.2 vezes; 7) aspecto singular nesta nova espécie é a presenca de um

sulco longitudinal no 3° tergito que separa a epipleura até o espirdculo. A

epipleura do 3° tergito separada por sulco longitudinal é uma das caracteristicas

da tribo Anomalonini; em Therionini, onde se inclui Philodrymus, a epipleura do

tergito III nao é separada por un sulco longitudinal.

Philodrymus e Podogaster que lhe é muito afim, sao dois pequenos generos

Neotropicais de Anomaloninae, com caracteristicas bastante especializadas princi-

palmente nas asas (veias longitudinais da 28 célula discoidal tendem a se unir

na parte basal, nimero de hamulos reduzido de 3 a 5, extremidade distal da

veia anal da asa anterior bem afastada da margem da asa e com mancha carac-

teristica que a une com a margen) e genitalia (edeago com o 4pice da parte

proximal geniculado e achatado) (Gauld, 1976:82), e que se distinguem entre

si principalmente pelo maior alongamento da projecdé6 posterior do propdéddeo e

maior convergéncia ventral dos olhos em Pologaster que em Philodrymus.

Philodrymus townesi se distancia do padrdé® genérico principalmente no alar-

gamento e alongamento posterior inferior da gena, na mudanga em arco da carena

genal que se une a base da mandibula, no grande alongamento do ovipositor e

no sulco longitudinal do 3° tergito, que separa a epipleura até o espirdaculo.

AGRADECIMENTOS. O autor agradece as profs. Yoshiko S. Kumiyoshi e

Bernadete L. de Oliveira a ajuda na obtengao dos frutos de cedro e cuidados com

os insetos obtidos em laboratério e aos profs. Albino M. Sakakibara e Pe. Jesus

S. Moure as fotografias e leitura do original.

REFERENCIAS

CRESSON, E. T. 1874. Descriptions of Mexican Ichneumonidae. Proc. Acad.

Nat. Sci. Philad. 1873:374-413.

GAULD, I. D. 1976. The classification of the Anomaloninae (Hymenoptera:

Ichneumonidae). Bull. Brit. Mus. Nat. Hist. (Ent.) 33(1):1-135.

SZEPLIGETI, G. V. 1906. Neue exotische Ichneumoniden. Ann. Hist. Nat.

Hung. 4:119-156. |

TOWNES, H. K. & TOWNES, M. 1966. A catalogue and reclassification of

Neotropic Ichneumonidae. Mem. Amer. Ent. Inst. 8:1-367.

TOWNES, H. K. 1971. Genera of Ichneumonidae. Mem. Amer. Ent. Inst.

Ly sie ore.

Graf: Ichneumofauna 401

Figs. 1-4. Philodrymus townesi, n. sp.: 1, Femea, em vista lateral

(holotipo). 2-4, cabeca do macho em vistas anterior, lateral e superior.

Fig. 5, Pupa de Hypsipyla grandella, em fruto (parte) de Cedrela fissilis.

INSECTOS QUE DESTRUYEN EL ESPARTILLO (STIPA BRACHYCHAETA)

Y SUS PARASITOIDES

Luis De Santis

Director del Museo de La Plata. Profesor Emérito

Marta S. Loiacono de Silva

Museo de Ia Plata. Miembro de la Carrera del Investigador Cientifico

(Consejo Nacional de Investigaciones Cientificas y Técnicas)

Abstract

The principal enemy of "espartillo" in Buenos Aires, La Pampa and Entre

Rios provinces, is a wasp belonging to the genus Jetrdmesa,a new species here

described as T. adrianae De Santis, providing also bionomic data on this species.

It's the first record of the genus Tetramesa (Eurytomidae) in Argentina. Less

importance have the Gramineae moth borer not utilizable for biologic control for

obvious reasons. 7.adrianae also is not a promissory in weed struggle: the

single form for use could be as follows: (1) by means of its propagation, rang-

ing the plague geographic distribution; (2) by means of its massive breeding

and liberation in order to increase the populations. This fact requires to build

special insectaries for that purpose, together with previous studies, and (3)

by means of an integrated control, together with other procedures.

Otherwise, Tetramesa has various natural enemies also studied in this

paper; they are: Mimencyrtus stipitatus De Santis, new genus and new species

(Pteromalidae); Eurytoma cordoi De Santis, new species (Eurytomidae); Merce-

tina entrerriana De Santis,: new species (Eupelmidae) and Horismenus seminiger

De Santis, new species and H. christenseni (Blanchard, 1942) both belonging to

the family Eulophidae.

The types of the new species and all other material studied in this paper

are deposited at Museo de La Plata collections.

Este trabajo ha sido llevado a cabo a fin de responder a consultas que nos

han formulado la doctora Adriana Rios de Saluso, de la Estacién Experimental

Agropecuaria de Parana (Entre Rios) dependiente del Instituto Nacional de Tec-

nologia Agropecuaria (INTA) y del ingeniero agrédnomo Hugo A. Cordo, del

Laboratorio de Control Biolégico de Malezas que el Departamento de Agricultura

de los Estados Unidos tiene instalado en Hurlingham (Buenos Aires). Parte de

los materiales estudiados nos fueron facilitados por el ingeniero agrénomo Cordo

y la doctora Rios de Saluso y el resto ha sido criado por nosotros en matas de

Stipa brachychaeta que nos han enviado, periéddicamente y a nuestro pedido, la

doctora Rios de Saluso y el licenciado Julio C. Ves Losada, de la Estacién Expe-

rimental Agropecuaria (INTA) de Anguil (La Pampa). Dejamos constancia, ademas,

de que su realizacién ha sido posible gracias a un subsidio que nos ha otorgado

la Comisién Administradora del Fondo de Promocién de la Tecnologia Agropecuaria

(CAFPTA) y en tal sentido, debemos destacar el amplio apoyo brindado por su

Presidente, el ingeniero agrénomo Rail H. Quintanilla.! A la profesora doctora

Elsa M. Zardini, de la Facultad de Ciencias Naturales y Museo de La Plata, le

debemos la importante informacién que damos sobre la planta huésped y al licen-

ciado Andrés Boltoskoy, una paciente traduccién de la obra de la doctora

M. D. Zerova que tantas veces citamos en este trabajo.

lPallecié en la ciudad de Cérdoba (Reptblica Argentina) el 20 de agosto

de 1981 (véase De Santis, 1981, Rev. Soc. ent. arg., 40 (1-4): 106, 248).

402

De Santis: Insectos destruyen espartillo , 403

Los tipos de las nuevas especies, como asi también el resto de los materiales

estudiados, han sido incorporados a las colecciones del Museo de La Plata.

LA PLANTA HUESPED

La Stipa brachychaeta Godron, 1853, conocida con los nombres vulgares de

pasto puna, pasto indio, paja vizcachera, paja india y espartillo, se la encuentra

en las provincias de Salta, Tucuman, Santiago del Estero, San Luis, Catamarca,

Cérdoba, Santa Fe, Entre Rios, Buenos Aires y La Pampa. Dicen Marzocca,

Marsico y del Puerto (1979) que "ha sido citada como maleza de algunos cultivos

del norte de la provincia de San Luis, de Santa Fe (especialmente en el departa-

mento de San Cristébal) y de Cérdoba, donde es una de las mds temibles malezas

de la alfalfa". Merced a sus semillas originadas de flores cleist6gamas, ocultas

en el suelo, reaparece en los campos, atin después de afios de labranza; es muy

resistente al pisoteo [véase Burkart (1969)]. Para informarse sobre esta planta,

en relacién con la bionomia de los insectos que estudiamos mds adelante, es im-

prescindible la consulta de los trabajos de Caro (1966) Cano y Eilberg (1969)

Caro y Sanchez (1971) y Eilberg (1974). Expresa esta iltima investigadora que

"las canas que fructifican apicalmente se secan y deterioran. Asi, es dable

hallar en matas de varios afios, cafias verdes, cafias secas y cafias secas de las

cuales sdélo queda la parte basal".

En relacién con la utilidad que prestan o el perjuicio que pueden llegar a

ocasionar los insectos que aqui estudiamos, es importante destacar que la

S. brachychaeta también puede comportarse como una planta &Gtil; en efecto, dice

Burkart (1969) que es "mala forrajera, pero que es un recurso en potreros sobre-

pastoreados y el rebrote tierno es aprovechado a falta de mejores pastos": al

respecto, hay un estudio quimico de la doctora Abiusso (1962) y otro sobre la

fibra por Guth (1961) y segin la ingeniera agrénoma Eilberg (1969) con los

frutos de origen cleist6gamo pueden hacerse "pochoclos" o pororé.

LOS INSECTOS QUE LA DESTRUYEN

En las muestras estudiadas hemos determinado que destruyen las plantas de

espartillo una especie de microhimendptero del género Tetramesa que es nueva

para la ciencia y la describimos més adelante y una oruga taladradora de las

cafias. También se han hallado larvas de dos especies de dipteros cecidémidos

y de un tisandéptero tubulifero pero que, al parecer, no revisten mayor impor-

tancia como enemigos de la planta. Nos ocuparemos a continuacién, del estudio

de todas estas especies empezando por el microhimenéptero por ser el que merece

mayor atencién como destructor del espartillo. Ahora bien, como es ésta la pri-

mera vez que se cita una especie de Tetramesa para la fauna argentina, antes

de estudiarla, daremos los datos correspondientes al género:

EURYTOMIDAE

Género TETRAMESA Walker

!sosoma Walker, 1832 (no Billberg, 1820). Ent. Mag. 1: 13-14,

Tetramesa Walker, 1848. List Spec. Hym. Ins.Brit.Mus. Chalc. 2:

104, 154,

_Harmolita Motschulsky, 1863. Bull. Soc. Imp. Nat. Moscou. 36 (2): 58.

Xanthosoma Ashmead, 1888. Entom. Amer, 4: 42-43,

Isosomocharis Ashmead, 1888, ibid, 4: 42-43.

Urios Girault, 1911. J. N. VY. Ent. Soc. 19: 175.

Exanthosoma Girault, 1915. Mem. Qd. Mus. 4: 265.

Isthmosoma Hedicke, 1921. Arch. f. Naturg. 86 A 11: 165,

Gahaniola Erdoes, 1952. Rovart. Kozlemény. (n. s.) 5: 117,

Antola Claridge, 1961. Trans. Soc. Brit. Ent. 14: 174.

Morfologia: Para informarse sobre morfologia externa y anatomfia, hay que

que recurrir al trabajo de James (1926) sobre T. hyalinipennis (Walker, 1832).

La estructura del sistema reproductor femenino en Tetramesa y otros euritémidos,

ha sido estudiada por Copland y King (1972).

404 Contrib. Amer. Ent. Inst., vol. 20, 1983 —

Distribucion geografica: Segin Burks (1971) las especies de 7 etramesa

estan presentes en todo el mundo pero para la regién neotropical sélo tenemos

anotadas T. grandis (Riley, 1884) para México y T. tritici (Fitch, 1859) para

Colombia y recordamos que en cierta oportunidad llegaron a nuestras manos

ejemplares de una especie no identificada del Brasil.

No tenemos elementos para decidir si la nueva especie que describimos a

continuacién es autéctona o introducida.

Bionomia: Existen tres trabajos que son fundamentales para el estudio de

la bionomia de las Tetramesa: Phillips (1920), Claridge (1961) y Zerova (1976)

el primero referido a las especies nearticas y los otros dos a las. palearticas pero

que traen informacion sobre las investigaciones efectuadas con anterioridad, al-

gunas de ellas sobre especies de otras regiones.

TETRAMESA ADRIANAE DE SANTIS, N. SP.

Huevo: El huevo depositado es, en otras especies de 7etramesa,.incoloro y

translacido. Nosotros sélo hemos tenido oportunidad de observar los huevos

ovaricos representados, con mucho aumento, en la figura 7; es parecido al de la

especie T. scheppigi (Schlechtendal, 1891) representada en la figura 115 del

trabajo por Zerova (1976); ofrece una corta prolongacién en el polo anterior mi-.

eropilar y un largo pedtinculo en el polo opuesto que el insecto utiliza para fijar

el huevo en la pared interna de la cafia. Recuérdese que 7. scheppigi una

especie paleartica, se desarrolla sobre Stipa pennatay S. sabulosa.

Larva desarrollada: Representada en las figuras 3, 8, 9, 10 y 12. Es alar-

gada, blanquecina, mide 4 mm. de largo y pueden contarse muy bien, 13 segmen-

tos; la cabeza con 2 pares de setas sensoriales bien desarrolladas; ademas, son

perfectamente visibles 9 pares de espiraculos: uno mesotoracico, otro metatora-

cico y 7 ubicados en los primeros 7 segmentos abdominales.

Pupa: Representada en la figura 13.

Hembra adulta: Castafio oscuro o negro. Cabeza, escapo, Apice del pedicelo,

anillo, pronoto, coxas anteriores y segundo artejo de los trocanteres, de color

leonado, ennegrecido en la zona ocelar, en casi toda la mitad posterior del pronoto

y en el artejo apical de los tarsos. Base de los fémures intermedios, rodillas,

apice de las tibias, espolones, los cuatro primeros artejos de los tarsos, nerva-

duras alares y vainas del oviscapto, amarillo.

Vértice reticulado-punteado; frente, cara y mejillas, rugoso, con rugas lon-

citudinales; sutura genal marcada nada mas que por un corto trecho por debajo

de los ojos. Dorso del t6érax y pleuras, reticulado, rugoso en la parte posterior

del mesoescudo y en el escutelo, parapsides y axilas; pronoto con punteado ape-

nas perceptible y muy disperso. Estructuras del propodeo representadas en la

figura 4, con impresiédn mediana ancha. Gaster brillante con reticulacién apenas

marcada,

Cabeza y alitronco con setas cortas, escamiformes y reclinadas; gaster mas

pestafioso a partir del urotergito V. Ojos con setas cortisimas y muy dispersas.

| Escrobas bien marcadas, no marginadas; cara convexa; occipucio excavado

en la regién del cuello; ocelos en triangulo obtusaéngulo, los posteriores a dos

didmetros de las érbitas internas correspondientes; lineas post-ocelar y ocelo-

ocular en la relacién siguiente: 9 : 3.5; mejillas un poco mas cortas que el dia-

metro longitudinal de los ojos; mandibulas anchas en el 4pice, confusamente tri-

dentadas; antenas insertas un poco por encima de la linea inferior de los ojos;

el escapo llega nada mas que hasta el plano del vértice y la maza es biarticulada,

sin trazas de divisién en el artejo apical; dimensiones de cada artejo:

De Santis: Insectos destruyen espartillo 405

Longitud Anchura Longitud Anchura

Reverecesoves 0.047 G08G” Viet saveusccvet OUTe 0.057

L sv ioe. abs 0,243 O O6O ML ee CER OR ~ 0.072 0.067

LL ahCbW S's Odes 0.101 06082) Vil ia 4 wxaen 7 De0G2 0.067

anillo: «ai edly: 0.016 0,034 9 Wildes \. ders 0.062 0.078

NS epee ee 0.093 01069. TReaiwed wae 0.114 0.078

LV ats tac ‘ea 0.072 0.052

Longitud y anchura del collar del pronoto en la relacién siguiente: 10 : 25;

mesoescudo un poco mas corto que el escutelo; propodeo y espiraculos tal como

puede verse en la figura 4. Longitud de las alas anteriores 1.943; anchura

maxima 0.762, con setas marginales muy cortas; longitud de las nervaduras sub-

marginal, marginal, postmarginal y estigmatica, en la relacién siguiente: 67 ; 19 :

10 : 11. Longitud de las alas posteriores 1.486;. anchura maxima 0.349; longi-

tud de las setas marginales mas largas 0.047.

Peciolo corto; géster un poco mas largo que la cabeza y el alitronco reunidos

(85 : 75); el urotergito IV es el mas largo; los 6 primeros con margen apical

recto; proyeccién de las vainas del oviscapto sub-igual a la longitud del uroter-

gito VI.

Longitud del cuerpo 3.2.

Macho: Se diferencia de la hembra por la conformacién de las antenas, el

propodeo con estructiras distintas y la impresidn mediana mas estrecha; el pe-

ciolo notablemente mas largo y el gaster mas pequefio y truncado en el 4pice

(figura 6).

Antenas conformadas tal como se ve en la figura 5; dimensiones de cada

artejo:

Longitud Anchura Longitud Anchura

aie done ee j 0.047 WiGoe Vereaet ees sd 0.160 0.052

Pay aie ie 0.243 Ole Ee eg 0.144 0.047

Wink hence 0.098 Oeae ONE Se eas 0.119 0.047

BRIO Vode: 0.013 O2DO. OVI ae thes a 0.083 0.047

by yar ee 0.207 GO So Nee ake Va Cae 0.088 0.040

i Ga eee 0.052

Peciolo reticulado. Longitud del cuerpo 1.8.

Variaciones: Es muy variable la intensidad de la coloracién oscura del in-

secto y es asi, como se pueden hallar ejemplares de color castafio.

Distribucién geografica: Buenos Aires, La Pampa y Entre Rios. Localidad

del tipo: Paranda, en la Ultima de las provincias mencionadas.

Bionomia: Los primeros adultos aparecen en setiembre y van en aumento

hasta alcanzar un maximo de emergencia en diciembre que es cuando también co-

mienzan a aparecer los adultos de los parasitoides que estudiamos mas adelante.

En enero se siguen obteniendo adultos de estos altimos; las Tetramesa en cambio

nacidas de huevos depositados por estos adultos, no se transforman y siguen al

estado de larvas maduras hasta la primavera siguiente. Vale decir, entonces,

que pasan el invierno en ese estado; es por todo esto, que afirma Zerova (1976)

que en muchas especies de Tetramesa de bionomia conocida, el 90 por ciento del

tiempo de duracién del ciclo evolutivo lo pasan al estado de larva. Es interesante

anotar aqui, que las especies paledrticas T.eximia (Giraud, 1863) T. angustipennis

(Walker, 1832) y T. brevicornis (Walker, 1832) invernan al estado de pupa. En

relacién con esta diapausa invernal, no se puede dejar de mencionar el hecho

extraordinario de que las larvas de la especie T.phyllostachitis (Gahan, 1922)

pueden permanecer en el interior de las cafias de bambi por espacio de 22 meses

y empupar en la estacién siguiente. Hacemos esta observacién, porque tenemos

en el Laboratorio trozos de cafias secas con larvas maduras de T.adrianae

recibidas en abril de 1980 que, a la fecha, enero de 1981, atin no se han

406 Contrib. Amer. Ent. Inst., vol. 20, 1983

transformado. Llevadas a la cémara de cria, sdlo se logré que empuparan algu-

nas y se transformaran en adulto, recién cuando se las sometiéd a temperaturas

de 35° C. de dia y 20° C. por las noches. Venimos realizando observaciones

sobre esta especie desde 1977 y debemos anotar que nunca hemos podido sor-

prender ejemplares en cépula o hembras efectuando la puesta de huevos. Dice

Zerova (1976) que algunas Tetramesa para lograr maduracién de las génadas

necesitan de una alimentacién suplementaria y recordemos a propdsito de esto,

que existen euritédmidos que tienen un régimen mixto: entoméfagos al principio

y después fitéfagos, una vez que han consumido la presa.

Repetimos que no hemos tenido oportunidad de hallar huevos depositados en

las cafias examinadas pero es evidente que 7. adranae produce una sola genera-

cidn anual; de huevos depositados en primavera nacen larvas que completan su

desarrollo en diciembre pero que recién empupan y se transforman en adulto, en

la primavera siguiente. También puede apreciarse que la hembra deposita varios

huevos en cada entrenudo ya que nos ha sido posible comprobar en las cafas

examinadas que, por lo general, presentan unas 10 celdas aisladas y dispuestas

en sentido longitudinal pero abarcando la circunferencia de la cafia (figuras 8 a

12). Segtan Zerova (1976) estas celdas son excavadas por las larvas de la.

Tetramesa y quedan protegidas por una pared que construyen con trocitos de

sustancias que toman del tallo; cada celda mide 4 mm de largo por 1 de ancho.

El ataque se produce en los entrenudos de la base de las cafias pero a cierta

distancia del suelo, es decir, algo alejadas de la region del cuello de la planta.

Como ocurre con otras especies de Tetramesa este ataque provoca una reaccién

del vegetal que se deforma en el entrenudo correspondiente, tal como puede

apreciarse en la figura 11.

Hasta el presente no hemos hallado larvas de los primeros estadios en las

canas examinadas. Dice Zerova (1976) que una vez depositados, los huevos

hacen eclosién entre los 5 y 10 dias subsiguientes; luego el desarrollo larval es

muy lento segiin hemos explicado en el lugar correspondiente. Anotamos de paso,

que en una larva que habia iniciado el proceso de pupacién, llevada a la camara

de cria a 35° C. de dia y 20° C. de noche, el periodo pupal se prolongé por

espacio de 14 dias; para las especies paledrticas de bionomia conocida, Zerova

(1976) anota para ese periodo una duracién que oscila entre 15 y 30 dias. Para

estudiar el comportamiento de las larvas y pupas de 7. adrianae y de los para-

sitoides que las atacan, descubrimos la celda por medio de un bisturi y, efectuada

la observacioOn, volvemos a cubrirla con el trocito de tallo extrafdo y lo fijamos

con cinta adhesiva; esto permite repetir la observacién tantas veces como sea

necesario. |

La transformaci6n en pupa tiene lugar en la misma celda (figura 13) y cuando

aparecen los adultos, estos perforan la cafia con sus mandibulas y hacen un pe-

quefio orificio de salida tal como puede apreciarse en la figura 11. —

Para que se tenga una idea sobre las poblaciones de la 7. adrianae la natu-

raleza del dano que producen y la intensidad del ataque, damos, a continuacién,

los resultados que se obtuvieron con el examen de 100 cafias provenientes de una

misma mata, de Parana, recibida en diciembre de 1979;. la observacién fue efec-

tuada en enero de 1980 y los datos correspondientes se refieren a un solo entre-

nudo:

Entrenudos no atacados por Jetramesa ..... 59

Entrenudos con ataque de TJetramesa ...... oy 41

En esos 41 trozos de cafas atacadas por Tetramesa_ se contaron 216 celdas,

104 conteniendo larvas maduras vivas, 4 con adultos muertos de 7. adrianae y

Eurytoma cordoi y 108 vacias, es decir que habian sido abandonadas por los

adultos.

Hemos anotado precedentemente, que trozos de cafia con larvas maduras de

T, adrianae_ de matas recibidas en abril de 1980, atin no habian empupado en

enero de 1981, lo cual se vuelve a repetir ahora, con las 104 larvas vivas pre-

De Santis: Insectos destruyen espartillo 407

sentes en los trozos de cajfia utilizados en este experimento; queda la duda sobre

Si esto puede producirse en condiciones naturales, es decir en 1a planta en pie,

sobre todo teniendo en cuenta que las larvas maduras también se hallan en las

cafias secas; si asi fuese tendriamos algo similar a lo que ocurre con la Tetra-

mesa del bambt (T. phyllostachitis).

Danos: El ataque de 7, adrignae provoca un debilitamiento de la cafia alte-

rando la buena circulacién y haciéndola propensa al vuelco aparte de los proce-

sos irritantes que llevan a la formacién de las cecidias observadas. El dafio me-

canico que produce el insecto con el oviscapto y el aparato bucal abre vias a la

penetracién de otros insectos y a4caros y de gérmenes patégenos. En ningin

caso pudimos comprobar el ataque a los 6rganos reproductores como lo hacen,

por ejemplo, las especies palearticas T.cylindrica (Schlechtendal, 1891) sobre

Stipa capillata y T. punctata Zerova, 1965, sobre Stipa Jessingiana, Habia mucho

interés en investigar este punto debido a que 1a doctora Rios de Saluso (in litt.)

nos habia manifestado que al observar el interior de los tallos donde viven las

larvas de T.adrianae habia notado la ausencia de semillas cleist6gamas; queda por

investigar si existe alguna accién indirecta con esas consecuencias. De cual-

quiera manera que sea, debemos manifestar que ninguna de las especies dafiinas

que estudiamos en este trabajo, es capaz, por si sola, de eliminar a plantas que,

como éstas, exhiben tanta vitalidad.

Otro hecho que debe destacarse es la especificidad de las Tetramesa; como

lo ha hecho notar Phillips (1936) es rarisimo que una misma especie de Tetramesa

se desarrolle en dos o mas plantas huéspedes, Por nuestra parte, hemos exten-

dido nuestras investigaciones a otras especies de Stipa de Buenos Aires y La

Pampa y no hemos hallado en ellas, los insectos que aqui estudiamos.

Observaciones: Esta nueva Tetramesa es diferente de las especies norte-

americanas y euroasiaticas que viven en las St/pas; la mds parecida es T. acicu-

lata (Schlechtendal, 1891) pero se diferencia por la coloracién y también, por la

conformacién de las antenas. Esta dedicada a su descubridora, la doctora Adria-

na Rios de Saluso. |

Materiales estudiados: 1 F holotipo, 1M alotipo y 41 F y 33M paratipos,

Parana (Entre Rios) 1979-1980, Saluso, leg. ;

LAS ORUGAS TALADRADORAS

En las muestras de espartillo recolectadas en La Pampa se determiné la pre-.

sencia de orugas taladradoras. Han sido estudiadas por la Licenciada Griselda

Armesto, quien nos informa que deben referirse al género Diatraea; se desarro-

flan en la parte inferior de las cafias. Desgraciadamente, no se pudieron obte-

ner los adultos por lo que queda pendiente la determinacién de las mismas.

En 1978, recibimos una muestra de espartillo procedente de Parana que al-

bergaba una crisalida pero en este caso tampoco pudimos obtener la mariposa

debido a que estaba parasitada por un himendédptero icneuménido, segin referi-

mos mas adelante. Es muy parecida también a las crisdlidas de Diatraea pero no

podemos afirmar que se trate de la misma especie que la que se obtuvo de mate-

riales procedentes de La Pampa.

LOS DIPTEROS CECIDOMIDOS

El ingeniero agrénomo Cordo hallé en cafias de espartillo recolectadas en

Campana (Buenos Aires) numerosas larvas de color anaranjado (figura 24) co-

rrespondientes a un diptero de la familia Cecidomyiidae. La hemos incluida: en

la subfamilia Cecidomyiinae, supertribu Cecydomyiidi tribu Contariniini y de

acuerdo con los datos que da Harris (1966) podria tratarse de una especie del

género Contarinia Rondani, 1860; la bibliografia respective menciona algunas

especies que viven en las espiguillas de las Gramineas. Queda pendiente su

clasifi¢acion definitiva para cuando pueda contarse con ejemplares adultos y

también el estudio de su comportamiento en relacidén con el espartillo para cuan-

do se disponga de suficiente cantidad de material vivo.

408 Contrib. Amer. Ent. Inst., vol. 20, 1983

Las muestras procedentes de Parand también nos proporcionaron materiales

de dipteros pertenecientes a esta familia: una larva que estudiamos en el capitulo

dedicado a los parasitoides y entomdfagos y otra de color anaranjado que se en-

contraba en una celda de Tetramesa, por debajo de una larva seca y, ademas,

un adulto hembra obtenido de materiales que se colocaron en la cAémara de cria;

se trata de un representante de la subfamilia Lestremiinae, tribu Catochini;

como es sabido, las larvas de las especies de esta iltima tribu se comportan como

sapréfagas o copréfagas, de manera que no ofrecen mayor interés de acuerdo con

las finalidades que se persiguen con esta investigacién.

Tampoco puede prestarse, por el momento, mayor atencién a la especie de

tisanoptero tubulifero presente en la iltima muestra legada de Parana en enero

de 1981, por dos razones: en primer lugar, por ser ésta la primera vez que se

la recibe y luego, porque si bien es cierto que existe un buen nimero de tisan-

opteros que se comportan como fitédfagos, son muchos los tubuliferos que son pre-

datores o se alimentan de hongos y algas.

UTILIZACION DE ESTOS INSECTOS EN LA LUCHA BIOLOGICA CONTRA

EL ESPARTILLO

De la exposicién que acabamos de hacer se infiere facilmente que, a lo que

hasta hoy se sabe, es 7. adFianae la tnica especie que podra ser utilizada contra

esta maleza pero, a juzgar por los datos que poseemos, el porcentaje de parasi-

toidismo natural es relativamente bajo con el agravante de que se esta frente a

una planta que exhibe una gran vitalidad y un extraordinario poder reproductor.

Ademas, segin se podra ver mas adelante, T. adrianae tiene, a su vez, varios

parasitoides que la destruyen. Por todo esto, la Gnica forma de utilizarla seria

poniendo en practica los procedimientos siguientes, todo.lo cual requiere una serie

de estudios previos y, en algunos casos, sa instalaciones correspondientes; son

estos:

1°)- Su propagacién, para tratar que la distribucién geografica de T. adria-

nae coincida con la que tiene la planta;

2°)- Su cria masal y luego su liberacién, a fin de incrementar las poblaciones

de este insecto Util, lo que requiere otros estudios previos y la instalacién de in-

sectarios con esa finalidad y

3°)- Su utilizacién combinada con otros of sdatcectehiteia' aie de lucha, es decir,

un control integrado de la plaga.

-LOS PARASITOIDES Y OTROS ENTOMOFAGOS

Estudiamos en este capitulo, los parasitoides y otros entomdéfagos que atacan

a los insectos tratados en paginas precedentes y mencionamos aquellos otros que

han sido hallados en las muestras sin que se haya podido establecer sus relacio-

nes con estos u otros insectos que viven en la Stipa brachychaeta,

LOS PARASITOIDES DE TETRAMESA ADRIANAE

Esta especie, la mas importante como enemigo natural del espartillo, es ata-

cada, a su vez, por varios entomdfagos, todo los cuales reducen, de manera

apreciable, la eficaz accién destructora que 7. adrianae ejerce sobre la plaga

que se desea combatir; los estudiamos a continuacidén:

EURYTOMIDAE

EURYTOMA CORDOI,N. SP.(FIGURAS 14, 17 y 20)

Larva madura: No hemos podido diferenciarla de la de 7. adrianae es pro-

bable, entonces, que alguna de las larvas que hemos atribuido a esta iltima es-

pecie, lo sean, en realidad, de E. cordo/.

Hembra adulta: Negro, con poco brillo. Tégulas de color castafio. Escapo,

apice del pedicelo, nervaduras alares y patas, excepto las coxas, amarillento,

irregularmente ennegrecido en la parte central de los fémures y tibias y en el

De Santis: Insectos destruyen espartillo 409

artejo apical de los tarsos. El resto del pedicelo y el flagelo de las antenas, par-

dusco. Alas hialinas.

Cabeza, propleuras, la mayor parte de las mesopleuras y las coxas, reticu-

lado. Pronoto, mesoescudo, escutelo, parapsides y axilas, reticulado-punteado,

con puntos setiferos poco profundos; propodeo rugoso con depresién mediana

profunda y bien marcada. Gaster débilmente reticulado. 3

Ojos pestafiosos, con setas cortas dispersas. Cabeza, altitronco, apéndices

y quinto apical del g4ster, con setas blanquecinas.

Cabeza vista de frente subtriangular, mas ancha que larga (34 : 29) escro-

bas bien excavadas, no marginadas; mejillas un poco mas cortas que el didmetro

longitudinal de los ojos (12 : 16) ocelos en triangulo obtusaéngulo; lineas post-

ocelar y ocelo-ocular en la relacién 9 : 4; mandibulas con dos dientes bien mar-

cados y una ancha truncadura oblicua interna; antenas insertas en el centro de

la cara, bien por encima de la linea inferior de los ojos; maza biarticulada, sin

trazas de divisiédn en el artejo apical; dimensiones de cada artejo:

Longitud Anchura Longitud Anchura

Loree em 0.036 0.026 Wales een sa et 0.062 0.047

TS ICE ORS ses 0.233 0.044 Md Las) « age thee 0.062 0.057

LY vackiport& teed 0.072 0.044 bok PSA ay 0.057 0.062

SND 46s: : 0.011 0.028 ATES Pe rey as 0.062 0.101

Es wcas £ 4 a8 0.062 0.036 Lt De eee ape 0 10k 0.101

IN eae hime ate 0.052 0.041

Longitud y anchura del collar del pronoto en la relacién 10 : 16, mas corto

que el mesoescudo que es, a su vez, mas largo que el escutelo; espirdculos elip-

ticos. Longitud de las alas anteriores 1.461; anchura maxima 0.648; longitud de

las setas marginales mas largas 0.021; longitud de las nervaduras submarginal

marginal, postmarginal y estigmatica en la relaci6n 51 : 15 : 15 : 10.5.

Peciolo muy corto; gaster un poco mas largo que la cabeza y el alitronco

reunidos; el urotergito IV es m4s largo que los tres primeros reunidos y también

que los siguientes tomados en conjunto; oviscapto apenas saliente.

Longitud del cuerpo 2.4.

Macho: Se diferencia de la hembra por la conformacién de las antenas, el

peciolo mas largo y el gdster mds pequefio; el tercero es el mas largo de los

urotergitos.

Antenas conformadas tal como se ve en la figura 20; dimensiones dr cada

artejo:

Longitud Anchura Longitud Anchura

Wik PG ow shake 0.036 0.028 W wtiacannisa <aee 0.098 0.014

bic erie ea OHS % 0.186 0.052 Rh « iad Utena wen 0.093 0.052

Bb scam Sota 0.062 0.041 Ad as hack gals 0.067 0.057

ONG i. caer 0.005 0.028 hat area Sues 0.067 0.057

he ee ee ee 0.119 0.052 Lheyidict eeaices 0.078 0.049

IV 4 < pike when 0.098 0.062

Peciolo el doble mas largo que ancho en la parte distal.

Longitud del cuerpo 1.5.

Distribucién geografica: Entre Rios. Localidad del tipo: Parana.

Bionomia: Parasitoide de T, adrianae,

Observaciones: Esta nueva especie puede ser comparada con E, pojincarei

Girault, 1913, del Paraguay; se diferencia por su tamafio menor y por la confor-

maci6én de las antenas. Esta dedicada al ingeniero agrénomo Hugo A. Cordo.

Materiales estudiados: 1 F holotipo, 1 M alotipo y 18 F y 70 M paratipos,

Parana (Entre Rios), 1979-1980, Saluso leg.

410 Gontribs: Amer. Ents insti; vol. 20, 1983:

El parasitoide que estudiamos a continuacién tiene que referirse a un género

cuya presencia en la Reptblica Argentina se senala aqui por primera vez, por

lo que damos los datos correspondientes al mismo. Por otra parte, también cree-

mos que es esta la primera vez que se describe el.macho.

EUPELMIDAE

Género MERCETINA Bolivar

MERCETINA BOLIVAR Y PIELTAIN, 1933: 205.

Macho: Macréptero. Escrobas bien excavadas; maza de las antenas entera.

Pronoto sin surco mediano, sin fosetas laterales en el collar y sin el par de pin-

celes de setas fuertes; surcos parapsidales ausentes, en su lugar un par de im-

presiones que divide al mesoescudo en tres lédbulos; escutelo convexo, brillante

en el A4pice. Tarsos intermedios normales; propodeo bien desarrollado, reticulado

y con quillas mediana y espiraculares; espiraculos elipticos, algo alejados del

borde anterior. Gdaster deprimido, de forma oval.

Morfologia: Este género tendra que ser interpretado con las rectificaciones

a la descripciOn original formuladas por Ferriere (1954).

MERCETINA ENTRERRIANA DE SANTIS, N. SP. (FIGURAS 15, 18, 21, 26 y 27)

Larva madura: Sinninguna seguridad, pensamos que puede corresponder

a esta especie, la larva ma@ura representada con mucho aumento en las figuras

26 y 27. :

Hembra adulta: Castafio oscuro, con reflejos verdosos, dorados y purpt-

reos en la cabeza, alitronco, gaster y en los fémures y antenas. Base y canto

inferior del escapo, trocdénteres medios y posteriores, espolones, los cuatro pri-

meros artejos de los tarsos intermedios, basitarsos posteriores y una mancha

central en las vainas del oviscapto, amarillo, en partes ennegrecido. Trocanteres

y rodillas de las patas anteriores, aclarecidos.

Frontovértice y cara con lineas estructurales transversales que se vuelven

longitudinales en las sienes, mejillas y parte inferior de la cara; occipucio liso

y brillante en la parte que rodea al foramen. Protdrax reticulado, liso en la parte

excavada del pronto y casi liso en la regién esternal. Mesoescudo, escutelo y

axilas, reticulado; mesopleuras con lineas estructurales concéntricas; metapleuras

y coxas también reticuladas. Gdster con reticulacién débilmente marcada.

Cabeza, alitronco, gaster y apéndices, con setas blanquecinas o grisaceas.

Ojos glabros. Pronoto con los dos pinceles caracteristicos de setas fuertes, eri-

zadas. Metanoto lateralmente y canto posterior de las metacoxas, con grupos

caracteristicos de setas blanquecinas. Setas apicales del escutelo también blan-

quecinas.

Cabeza tan ancha como el toérax, convexa); vista de frente de contorno casi

circular, poco mas larga que ancha; frontovértice ancho; escrobas poco excavadas,

no marginadas; mejillas tan largas como la mitad del diaémetro longitudinal de los

ojos; sutura genal presente; ojos medianos; ocelos en tridngulo equilatero; lineas

-post-ocelar y ocelo-ocular en la relacién 5 : 2; mandibulas tridentadas; palpos

maxilares de 4 artejos, el artejo apical one mas largo que los demas; labiales

de 3; antenas insertas a la altura de la linea inferior de los ojos; dimensiones

de cada artejo:

Longitud Anchura Longitud Anchura

Buat eee 0.057 0,044 Mite. Co ea et Res 0.065

Pi nee ewe 0.330 0.052 Vila maces ues oa PGR? 0.067

Peake eo ek 0.109 0.052 MU Ee seine aw 0.067 0.075

UAIELO 9 in sae 0.036 0.036 LS ae ee eee 0.072 0.078

RE eke als asa yc 0.088 0.041 bee einer 0.098 0.093

OVS eaees ss 0.098 0.047 2 ER PR ah Sa 0.062 0.093

Ws cae ONY 0.103 0.059 i ards ihe Te as 0.041 0.072

De Santis: Insectos destruyen espartillo 411

Pronoto de igual longitud y anchura. Alas subtriangulares, truncadas en

el Apice y con una nervadura longitudinal central que lleva 7 setas bastantes

largas; longitud 0.160 y ancho 0.109. Patas anteriores con fémures engrosados

y tarsos largos, mas largos que las tibias correspondientes; fémures y tibias in-

termedios largos; estas tlitmas llevan 5 espinas gruesas y oscuras en el 4pice;

su espolén es grueso y tan largo como el basitarso respectivo; este Gltimo con un

par de espinas gruesas y oscuras en el 4pice, lo mismo que el segundo tarsito;

ademas, los cuatro primeros tarsitos llevan en la parte inferior la doble hilera de

espinas claras caracteristicas del género. Patas posteriores con los fémures algo

abultados y las tibias un poco mas largas que los tarsos correspondientes.

Gdster ovoideo, voluminoso, excluida la proyeccién del oviscapto, tiene la

misma longitud del alitronco; los 4 primeros urotergitos ligeramente escotados en

el margen apical y los dos ultimos muy cortos; placas cereales ubicadas muy cerca

del apice; proyeccién del oviscapto igual a dos quintos de la longitud total del

gaster.

Longitud del cuerpo, excluida la proyeccién del oviseapto, 2.5.

Macho: Cabeza y alitronco verde con reflejos purpireos en la cabeza y en

la parte anterior del pronoto. Radicula, pedicelo y flagelo de las antenas, patas

y gaster, de color castafio oscuro, este iltimo con reflejos verdes en el primer

urotergito y purptireos en el resto. Apice del pedicelo y el escapo, de color

castafio-amarillento. Trocanteres, rodillas y Apice de las tibias intermedias, sus

espolonos y los dos primeros artejos tarsales, casi toda la mitad basal de las

tibias posteriores y sus espolones y basitarsos, blanco.1! Alas subhialinas con

nervaduras parduscas.

Dorso del téraz reticulado. Gdaster con reticulacién mds débil.

Antenas conformadas tal como se ve en la figura 21; dimensiones de cada

artejo:

Longitud Anchura . Longitud Anchura

Oe cicaia ececncg lates 0.036 De | oc Wie 5 a gig asks 0.057 0.039

tT cbtese < oannae 0.222 DD 4O: 6 Mis iccecaty sind 0.052 0.041

BE babies «ue eo 0.093 US ila Je 8 ie eaten 0.052 0.047

anillo. , 0.026 02006) 6 VA aaiewns 0.047 0.047

BD ss avs 0.047 oO es Oe aay 0.047 0.054

Rey oe 0.057 Ce OR eran we 0.156 0.054

Vieeiig ox al cae 0.057 0.039

Longitud de las alas anteriores (figura 18) 0.965; anchura maxima 0.381;

longitud de las setas marginales mas largas 0.036; longitud de las nervaduras

Submarginal, marginal postmarginal y estigmatica, en la relacién siguiente:

25 : 20 : 14: 9. Longitud de las alas posteriores 0.775; anchura maxima 0.121;

longitud de las setas marginales mas largas 0.047.

Longitud del cuerpo 1.2.

Distribucién geograéfica: Entre Rios. Localidad del tipo: Parand.

Bionomia: Parasitoide de T, adrianae,

Observaciones: Utilizando la clave de Ferriere (1954) se llega a

Bolivar, 1933, que es la especie tipo del género pero que se diferencia de

ella por su tamafio menor, la cabeza mas carta y el escutelo sin el mechén de

setas negras; tambien es diferente la conformacién de las antenas.

INo doy los datos correspondientes a las patas anteriores porque estan

cubiertas por la goma y no se distinguen bien los colores.

412 Contrib. Amer. Ent. Inst., vol. 20, 1983

Materiales estudiados: 1 F holotipo, 1 M alotipo y 14 F paratipos, Parana

(Entre Rios) 1979-1980, Saluso leg.

PTEROMALIDAE

Género MIMENCYRTUS De Santis, nov.

Hembra: Cabeza vista de frente subtrapezoidal, mas ancha que larga;

cara convexa; frontovértice ancho; clipeo cuadrado, mal delimitado, con margen

apical recto o ligeramente cdéncavo; longitud de las mejillas igual a la mitad del

diadmetro longitudinal de los ojos; una mandibula con 2 dientes y una truncadura

oblicua interna y otra con un diente medio y dos truncaduras; ojos medianos;

zona ocelar levantada; ocelos en tridngulo obtusangulo, los posteriores a 2 dia-

metros del barde del occipucio.ya algo mas de las Orbitas internas correspondien-

tes; antenas con la conformacion 1, 1, 3, 5, 3, insertas bien por encima de la

linea inferior de los ojos; distancia de los puntos de insercién al borde apical

del clipeo el doble de la que media al ocelo anterior.

Alitronco, incluyendo las coxas, casi completamente reticulado; mesopleuras

con zonas lisas por debajo de la insercién de las alas. Collar del pronoto con

borde redondeado; surcos parapsidales incompletos, marcados nada mas que en.

la mitad anterior; escutelo poco convexo; propodeo corto, con quilla mediana y

pliegues laterales; espiraculos pequefios, elipticos, algo salientes y bastante ale-

jados del borde anterior; alas anteriores con specu/um basal amplio y con setas

discales muy cortas; nervadura marginal con engrosamiento en la mitad basal muy

aparente. Patas fuertes; espolén de las tibias intermedias largo y fino, casi tan

largo como el basitarso correspondiente; tibias posteriores con 2 espolones bien

diferenciados.

Peciolo cortisimo; gaster mds ancho que el alitronco y mas largo que éste

y la cabeza reunidos, aplanado en el dorso; el urotergito I es el mas largo y II

y VII los mas cortos, los restantes subiguales; oviscapto poco saliente.

Macho: Muy diferente de la hembra por la conformacion de las antenas,

el propodeo comparativamente mas largo lo mismo que el peciolo y el gaster mas

corto y mas estrecho. Anillos laminares; artejos del flagelo cilindricos, alargados

con numerosas sensorias longitudinales en 2 0 3 series entremezcladas; el artejo

apical subcénico. Los urotergitos II, VI y VII son los mas largos.

ESPECIE-TIPO: MIMENCYRTUS STIPITATUS DE SANTIS, N. SP.

Observaciones: Siguiendo las claves de Delucchi (1956) y Graham (1969)

se llega al género /schyroptix Delucchi, 1956, en cambio, si se emplea la que

han publicado Delucchi y Steffan (1956) tiene que ubicarselo cerca de Gugolzia

Delucchi et Steffan, 1956; este nuevo género se diferencia del primero, por la

insercién mas alta de las antenas y la amplitud del speculum: y de Gugolzia por

el tamajio del clipeo, las mejillas mas cortas, los artejos del flagelo tambien mas

cortos y la dentici6n mandibular distinta.

MIMENCYRTUS STIPITATUS DE SANTIS, N. SP.

Hembra: Verde, con reflejos purptreos en los fémures, coxas posteriores,

propodeo y gaster. Escapo, primer anillo, maza de las antenas, tégulas, nerva-

duras alares, trocanteres, rodillas, tibias y tarsos, amarillo, ennegrecido en el

artejo apical de los tarsos.

Cabeza reticulado-punteada, con puntos setiferos poco profundos y disper-

sos, en la cara. Gdaster con reticulaciOn mas débil.

Ojos glabros. Cabeza y toérax con setas blanquecinas, escamiformes y recli-

nadas; propodeo y gaster con setas mas largas, también blanquecinas.

El escapo de las antenas sobrepasa el plano del vértice; dimensiones de

cada artejo:

De Santis: Insectos destruyen espartillo 413

Longitud Anchura Longitud Anchura

iuais Gea beh 0.052 OO81 OLY ix eee Ooo 0g 0.103 0.057

tie Bee keen 0.292 ONT i Mietintwe ta Ke 64 0.093 0.062

LECe cin ea dees 0.083 OAT COPD cence bee 0.093 0.062

atlas Pe ve<s 0.010 OOSl Win aed es 0.083 0.067

E » ee 0.013 0.0384. VIllvveaesss 0.098 0.083

" + ee iar 0.016 Ooo. <TR sates ee 0.031 0.083

Lids «teks 205% 0.119 OV ORS codes « ve gin re 0.052 0.057

Longitud de las alas anteriores 1.676; anchura maxima 0.635; longitud de

las setas marginales mas largas 0.021; longitud de las nervaduras submarginal,

marginal, postmarginal y estigmatica, en la relacién siguiente: 46 : 28: 19: 14.

Longitud de las alas posteriores 1.334; anchura maxima 0.279; longitud de las

setas marginales mas largas 0.089. 7

Longitud del cuerpo 2.2.

Macho: GdAdster con una mancha blanca sub-basal. Funiculo de color cas-

tafio y la maza mas o menos ennegrecida en la mitad basal; longitud de cada

artejo: R, 0.047; I, 0.217; II, 0.047; ITI, 0.145; IV, 0.140; V, 0.140; VI, 0.129;

VII, 0.119; VIII, 0.103; IX, 0.067 y X, 0.062. Longitud del cuerpo 1.5.

Distribucién geografica: Entre Rios. Localidad del tipo: Parana.

Bionomia: Parasitoide de

Materiales estudiados: 1 F holotipo, 1M alotipo y 10 F y 6 M paratipos,

Parana (Entre Rios) 1979-1980, Saluso leg.

EULOPHIDAE

HORISMENUS SEMINIGER DE SANTIS, N. SP. (FIGURAS 16 y 19)

Hembra: Negro, con reflejos verdosos intensos en la cabeza, escapo de

las antenas, alitronco y en la coxas y fémures. Gdster con reflejos del mismo

color pero menos intensos. Rodillas, tercio 0 mitad apical de las tibias y los 3

primeros artejos de los tarsos, blanco. Alas hialinas con nervaduras parduscas.

Cabeza reticulada, con pequefios puntos setiferos bordeando las dérbitas

internas y en el clipeo; vértice reticulado-punteado. Pronoto, mesoescudo, pa-

rapsides, axilas y escutelo, reticulados; surco mediano del escutelo obliterado

en el tercio posterior; surcos laterales marcados por una serie de puntos; propo-

deo liso y brillante excepto en 2 bandas longitudinales centrales y en otras 2

mas cortas ubicadas a los costados de éstas en el tercio posterior, donde aparece

con punteado denso; el propodeo esta separado del metanoto por una depresién

transversa bastante profunda. Peciolo reticulado; primer urotergito liso, excepto

en una estrecha banda sub-apical, donde aparece reticulado; los restantes tam-

bién reticulados excepto a lo largo del margen apical.

Ojos pestafiosos. Gaster también bastante pestafioso, con setas mas bien

largas, lateralmente y en la mitad apical.

Cabeza un poco mas ancha que el térax, vista de frente subtriangular,

casi de igual longitud y anchura (26 : 28) escrobas excavadas, con quilla inte-

rantenal; ojos muy prolongados ventralmente y escotados a la altura de la in-

sercién de las antenas; mejillas muy cortas; insercién de las antenas muy baja

aunque un poco por encima de la linea inferior de los ojos; ocelos en tridngulo

apenas obtusangulo; lineas post-ocelar y ocelo-ocular en la relacién 7 : 3, los

posteriores un poco mas cerca del borde del occipucio que de las érbitas inter-

nas correspondientes; dimensiones de cada uno de los artejos de las antenas:

414 Contrib. Amer. Ent. Inst., vol. 20, 1983

Longitud Anchura Longitud Anchura

Bovests ese es 0.021 O02 FEV oes eee 0.057 0.047

Lope as ae 0.212 Os039 Ves cass ees 0.052 0.062

Pec ie ks 0.078 OG VE acces ts 0.062 0.072

Tes. ca es 0.067 02099 Wee eee 0.041 0.046

Longitud del proceso apical 0.021.

Collar del pronoto marginado. Longitud de las alas anteriores 1.295; anchu-

ra maxima 0.508; longitud de las setas marginales mas largas 0.052; longitud de

las nervaduras submarginal, marginal, postmarginal y estigmatica, en la relacién

siguiente: 25 : 46 : 3: 4. Longitud de las alas posteriores 1.029; anchura

maxima 0.254; longitud de las setas marginales mas largas 0.076.

Peciolo grueso, transverso. Gaster tan ancho como la cabeza y, aproxima-

damente, de igual longitud que el alitronco; el primer urotegrito es el mas largo

y ocupa la mitad de la longitud total del gaster; los siguientes, hasta el quinto,

gradualmente mas cortos; el sexto un poco mas:‘largo que el quinto y los 2 res-

tantes muy cortos; oviscapto apenas saliente.

Longitud del cuerpo 2.

Macho: Cara y frontovértice con reflejos purptreos; reflejos verdosos de

la base del gaster mas intensos. Antenas conformadas tal como se ve en la fi-

gura 19; dimensiones de cada artejo:

Longitud Anchura : Longitud Anchura

Sa ee 0.180 O.049: 2. Ve ies ewes 0.067 0.054

| es ee 0.059 OSG iN Liesis oa ses 0.067 0.054

ese 0.072 Oe04T Vil. cakes 0.098 0.047

TV Se 5 0.062 0.054

Longitud del proceso apical 0.021.

Peciolo subcilindrico ; relacién entre la longitud y su anchura en la base

12+. 40.

Longitud del cuerpo 1.5.

Distribucién geografica: Entre Rios. Localidad del tipo: Parana.

Bionomia: Criado de materiales de 7. adrianae. las especies de Horisme-

nus de bionomia conocida se comportan como endoparasitoides primarios de larvas

y pupas de lepidépteros, coleépteros, himendépteros y dipteros, a menudo con

tendencia al hiperparasitoidismo, de modo que queda por investigar, tanto para

ésta como para la especie siguiente, sus verdaderas relaciones con 7. adrianae

o otros insectos presentes en las muestras. |

Observaciones: Esta nueva especie tiene que incluirse en el grupo //x/vo-

rus caracterizado por presentar el escapo de las antenas oscuro y las tibias tam-

bién més o menos oscuras excepto en la base y en la parte apical; se ubica cerca

de H. hypatia (Girault, 1915) de Chile, de la que se diferencia por la coloracién

y el surco mediano del escutelo mas extendido.

Materiales estudiados: 1 F holotipo, 1M alotipo y 4 F y 4M paratipos,

Parana (Entre Rios) 1979-1980, Saluso leg.

HORISMENUS CHRISTENSEN! (BLANCHARD)

Holcopeltomorvha christenseni, Blanchard, 1942; 126.

Distribuci6én geografica: Cordoba y Entre Rios.

Bionomia: En Cérdoba ha sido criada de materiales. Alabama argillacea

(Hiibn.) y en Entre Rios, de materiales de 7. adrianae.

Observaciones: Este especie ha sido determinada por comparaci6én con ejem-

plares sintipos existentes en las colecciones del Museo de La Plata.

De Santis: Insectos destruyen espartillo 415

Material estudiado: 1 F Parana (Entre Rios) 1979-1980, Saluso leg.

- LOS PARASITOIDES DE LAS ORUGAS TALADRADORAS

De una pupa recolectada en Parana, se obtuvo el siguiente parasitoide:

ICHNEUMONIDAE

COCCYGOMINUS RUFIPES (BRULLE)

Pimpla rufipes Brullé in Lepeletier, 1846, Hist. Nat. Ins. Hym., 4: 102.

Distribucién geograéfica: BRASIL: Minas Gerais y Sao Paulo. REPUBLICA

ARGENTINA: Buenos Aires y Entre Rios.

Bionomia: Criada deP/usia sp. y de la pupa de un lepidéptero taladrador

no identificado.

Observaciones: Para mayor informacién sobre esta especie, deben consul-

tarse los trabajos de De Santis y Millan (1968) y, principalmente, el de Porter

(1970).

Material estudiado: 1 F Paranda (Entre Rios) 1-XII-1978, Saluso leg.

OTROS PARASITOIDES Y ENTOMOFAGOS

En la cémara de cria se han obtenido los 2 entoméfagos que mencionamos a

continuacién pero sin que se haya podido determinar sus relaciones con los demas

insectos estudiados en este trabajo.

SCELIONIDAE

MACROTELEIA LARGA MUESEBECK,

Macroteleia larga Muesebeck, 1977: 25.

Distribucién geografica: ECUADOR, BRASIL y REPUBLICA ARGENTINA:

Entre Rios.

Bionomia: Desconocida. Expresa Muesebeck (1977) que lo poco que se co-

noce acerca de la bionomia de las especies de este género, indica que se desa-

rrollan en los huevos de los ortépteros tetigonioideos.

Material estudiado: 1 F Parand (Entre Rios) 12 - II - 1978, Saluso leg.

CARABIDAE

SELENOPHORUS PUNCTULATUS (DEJEAN)

Harpalus punctulatus Dejean, 1829. Spécies général des coléoptéres, pag. 91.

Distribucién geografica: Rio Negro, Buenos Aires y Entre Rios.

Bionomia: Mencionamos este caraébido hallado en muestras procedentes de

Parana, porque muchas de las especies de bionomia conocida de esta familia,

estan sefialadas como activas cazadoras de orugas. Desconocemos si tiene alguna

relacién con los insectos que aqui estudiamos.

Observaciones: Determinada por comparacién con ejemplares de las colec-

ciones del Museo de La Plata.

CECIDOMYIIDAE

En setiembre de 1979, en una mata procedente de Parana (Entre Rios) se

hall6é la larva representada en la figura 23. Ha sido estudiada por la licenciada

Emma B. Lagrange, quien nos ha hecho saber que se trata de la larva de un

diptero cecidémido de la subfamilia Cecidomyiinae, supertribu Cecidomyiid/. tribu

Lestodiplosini, Como es sabido, los representantes de esta tribu son predatores

y se alimentan de acaros y de las larvas de otros cecidémidos.

416 Contrib. Amer. Ent. Inst., vol. 20, 1983

BIBLIOGRAFIA

ABIUSSO, N. G. 1962. Composicién quimica y valor alimenticio de algunas

plantas indigenas y cultivadas en la Reptblica Argentina. Rev. Invest.

agric. 16 (2): 93-247.

BLANCHARD, E. E. 1942. Pardsitos de Alabama argillacea Hbn. en la Reptblica

Argentina. Estudio preliminar. Ann. Soc. Cient. Arg. 134: 54-128.

BOLIVAR Y PIELTAIN, C. 1933. Estudios de algunos Eupélmidos nuevos de

Espana (Hym. Chalc.). Eos, Rev. Esp. Ent. 9 (3-4): 195-209. |

BURKART, A. 1969. Flora ilustrada de Entre Rios. Col. Cient. INTA 6 (2):

148-151. —

BURKS, B. D. 1971. A synopsis of the genera of the family Eurytomidae (Hy-

menoptera: Chalcidoidea). Trans. Amer. Ent. Soc. 97: 1-89.

CANO, E. & EILBERG, Blanca A. de. 1969. Morfologia de las flores cleisté-

gamas axilares de los nudos basales de Stipa brachychaeta Godr. Bol. Soc.

Arg. Bot. 11 (4): 295-298,

CARO, J. A. 1966. Las especies de Stipa (Gramineae) de la regién central

Argentina. Kurtziana 3: 7-119.

CARO, J. A. & SANCHEZ, E. 1971. Stipa brachychaeta, S. caudata y S. ber-

trandii. Darwiniana 16 (3-4): 638-643.

COPLAND, M..J. W. & KING, P. E. 1972. The structure of the female repro-

ductive system in the Eurytomidae (Chalcidoidea: Hymenoptera). J. Zool.

Lond. 166: 185-212. 3 :

DELUCCHI, V. 1956. Beitrage zur Kenntnis der Pteromaliden (Hym. Chalci-

doidea). Zeitchr. Angew. Entom. 39 (2); 229-257.

DELUCCHI, V. & STEFFAN, J. R. 1956. Gugolzia harmolitae, nouveau genre

et nouvelle spece, parasite de Harmolita romana. Bull. Soc. Ent. Fr. 61:

30-34.

De SANTIS, L. & MILLAN, Eulalia. 1968. Adiciones a la fauna argentina de

Icneuménidos (Hymenoptera). Notas Coms. Invest. Cient. Buenos Aires

Se ag eC

EILBERG, Blanca A. de. 1969. Rosetas de cleistégenos axilares de dos espe-

cies de Stipa.. Bol.- Soc. Arg. Bot. 11 (4): 303-305.

EILBERG, Blanca A. de 1974. Los granos del "pasto puna" (Stipa brachy-

chaeta Godr.) Su heterogénea morfologia. Idia. pags. 34-37. :

FERRIERE, Ch. 1954. Eupelmides brachypteres (Hym. Chalcidoidea). Mitteil.

schweiz. Ent. Gesel. 27 (1): 1-21.

GRAHAM, M. W. R. de V. 1969. The Pteromalidae of North-western Europe

(Hymenoptera: Chalcidoidea). Bull. Brit. Mus. Nat. Hist..(Ent.) Suppl.

16: 1-908. |

GUTH, Edith B, de 1961. Extraeccién y estudio de las fibras de "paja vizca-

chera" Stipa brachychaeta Godr. Rev. Invest. Agric. 15 (4): 619-639.

HARRIS, K. M. 1966. Gall midge genera of economic importance (Diptera:

Cecidomyiidae). Part 1: Introduction and subfamily Cecidomyiinae; super-

tribe Cecidomyiidi. Trans. R. Ent. Soc. Lond. 118 (10): 313-358.

JAMES, H. C. 1926. The anatomy of a British phytophagous Chalcidoid of the

genus Harmolita (Isosoma). Proc. Gen. Meet. Sci. Buss. Zool. Soc. Lond.

pags. 75-182.

MARZOCCA, A.; MARSICO, O. J. & del PUERTO, O. 1979. Manual de malezas.

Edit. Hemisferio Sur. pags. 202-203.

MUESEBECK, C. F. W. 1977. The parasitic wasps of the genus Macroteleia

Westwood of the world (Hymenoptera, Proctotrupoidea, Scelionidae). Techn.

Bull.oWis.; Dept. Agr. (L065) ¢.1-57 ; :

PHILLIPS, W. J. 1920. Studies on the life history and habits of the joint-worm

flies of the genus Harmolita (Isosoma) with recommendations for control.

Bull. U.S. Dept. Agr, (Profess. Pap. 808): 1-27.

PHILLIPS, 1936. A second revision of the chalcid flies of the genera Harmolita

De Santis: Insectos destruyen espartillo 417

(Isosoma) of America North of Mexico, with descriptions of 20 new species.

Tech. Bull. U. S. Dept. Agr. (518): 1-25.

PHILLIPS, W. J. & EMERY, W. T. 1919. A revision of the chalcidflies of the

genus Harmolia of America North of Mexico. Proc. U. S. Natl. Mus. 55:

433-471.

PORTER, Ch. C. 1970. A revision of the South American species of Coccygo-

minus (Hym. Ichneumonidae). Stud. Ent. 13 (1-4): 1-192.

ZEROVA, M. D. 1976. Fauna de la URSS. Himendépteros, VII (6) Calcidoideos

Fam. Eurytomidae, Subfamilias Rileyinae y Harmolitinae. Akad. Nauk. SSSR.

Zool. Inst. (N.S.) 110: 1-230 (En ucraniano). ,

DEDICATORIA

Dedico este trabajo que he realizado con la colaboraci6én de la Doctora Marta

S. Loiacono de Silva, al Doctor Henry K. Townes con motivo de cumplir, en la

fecha, 70 afios de edad. La hago con todo el respeto y admiraciédn que se mere-

ce la magnifica labor que ha llevado a cabo en un periodo que abarca, asi, media

centuria. Su voluminosa contribucién a la taxinomia de los himendpteros de la

Serie Parasitica y especialmente de los Ichneumonidae, es apreciada en todo el

mundo y esta considerada como basica para el estudio de dichos insectos pero,

como ya lo manifestara en otra oportunidad, falta, para coronar tan extraordina-

ria producci6n, que el Doctor Townes nos brinde el volumen 5 de su Genera of

Ichneumonidae que habra de tratar sobre los Ichneumoninae; pondria asi un

broche de. oro a tan brillante carrera.

En circunstancias tan emocionantes como esta, no puedo dejar de traer aqui

el grato recuerdo de su ejemplar esposa que lo ha acompafiado siempre y lo ha

apoyado en todo momento; una labor como la que ha llevado a cabo el Doctor

Townes no puede ser realizada sino se cuenta con el aliento de una compafiera

cabal y de la calidad humana con que siempre se ha prodigado la bondadosa

Marjorie. |

iMis mejores deseos para ambos!

La Plata, 20 de enero de 1982.

(firmado)

Ing. Agr. Dr. Luis De Santis

Director

Museo de La Plata

418 Contrib. Amer: Ent:sinst., vol, 20, 1983

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Fig. 1. Tetramesa adrianae, n. sp. Female.

De Santis: Insectos destruyen espartillo 419

Figs. 2-7. Tetramesa adrianae, n. sp.: 2, genitalia del

macho. 3, larva madura. 4, propodeo d la hembra. 5, antena

del macho. 6, propodeo, peciolo y gaster del mismo.

7, huevo ovarico.

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De Santis: Insectos destruyen espartillo 421

Figs. 11-13. Tetramesa adrianae, n. sp. 11-12, cafias

de espartillo mostrando el dafio causado por la larva.

13, pupa ocupando una celda.

Contriba Ameri: Ent: Inst..-ved.. 20; 1983

422

Female.

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De Santis: Insectos destruyen espartillo 423

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De Santis: Insectos destruyen espartillo 425

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18, ala anterior del macho de Mercetina entrerriana. 19, ant-

ena del macho de Horismenus seminiger. 20, antena del macho

de Eurytoma cordoi.

ay ge

antena del macho de Mercetina entrerriana.

1983

Contrib. Amer: Ent. lnst.,; vols 20,

426

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De Santis: Insectos destruyen espartillo 427

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sint..”.24, larva.ceé. dLptero. cecidomido Conteariniinix 25) oruga

taladradora. 26, larva de calcidoidea hallada oeldas.

2/7, detalle de la cabeza de la misma.

MALAISE TRAPS: THE TOWNES MODEL CATCHES MORE INSECTS

Robert W. Matthews and Janice R. Matthews

Department of Entomology, University of Georgia

Athens, GA 30602

ABSTRACT

Malaise traps of the Townes 1972 design captured 10 times as

many insects as a commercial Cornell model during a four week test

near Albany, New York in July 1977. Small changes in trap location

resulted in significant differences in capture efficiency.

INTRODUCTION

In 1962, Henry Townes published the construction plans for a

Malaise trap for capturing flying insects (Townes, 1962). Traps of

this design were subsequently used successfully the world over in

diverse situations, and his paper did much to attract the attention

of entomologists to the potential of the trap as a sampling and

collecting device. Ever working to improve trap collecting

efficiency, Townes worked out a new pattern for a much lighter,

more efficient trap that was easier to construct (Townes, 1972).

Many uses of the Malaise trap or variations of it have appeared in

the literature, some for rather specialized situations or sampling

protocols (see Steyskal, 1981).

Various authors have stressed the importance of Malaise trap

Placement (e.g. Gressitt and Gressitt, 1962; Matthews and Matthews,

1970, 1971), but Townes, perhaps more than anyone, was aware of the

influence of trap design on collecting efficiency. As he stated

(Townes, 1972) " ..there is much difference between a good design

and a poor one. A poorly designed trap may catch only a tenth of

the catch of a better one of the same size, but its inferiority may

be unsuspected until it is directly compared with the better trap."

Over the years, we have used a number of homemade and

commercially produced Malaise traps. Always, our subjective

impression was that the Townes traps were superior. However, to

our knowledge no comparisons of the relative collection efficiency

of alternative trap designs have been published. In summer 1977,

we had the opportunity to directly compare the 1972 Townes trap

with a commercially produced trap. We here summarize the results

of this study. It is a particular pleasure to dedicate this paper

to Henry Townes on the occasion of his 70th birthday. Through the

years a substantial proportion of the parasitic wasps in research

collections have been obtained through the use of Malaise traps,

and Henry Townes, by his enthusiasm for these traps, has done more

than anyone else to improve and popularize their use.

Matthews and Matthews: Malaise Trap A429

Figure 1. The two Malaise traps used in this study. Left,

the nylon khaki-colored, commercially produced Cornell trap, with

openings in four directions. Right, the bidirectional Townes 1972

trap made of dacron marquisette. Corner posts are 140 cm tall.

METHODS

For four weeks during July 1977 (July 1-28) we operated

Malaise traps at two locations in the E. N. Huyck Preserve,

Rensselaerville, New York. The research reported here was part of

a larger study comparing changes in insect fauna over a decade.

Thus, traps were set up in the identical spots where Traps 1 and 2

were located in 1967 (Table 1, Matthews and Matthews, 1970). At

each location, a Townes 1972 model trap was placed beside a

commercially available Cornell-type trap.1 Both are non-attractant,

interceptive devices that capture free-flying insects. They differ

in shape, arrangement of openings, fabric, and color (Fig. 1).

The Townes trap we constructed was 10% smaller than the directions

specified, but the materials and coloration were faithfully

followed. The Cornell trap collected from four directions,

Sampling a total air space of 4.0 m2 (1.0 m2 per opening). Our

Townes trap sampled an air space of 1.5 m2 from each of two

directions. Thus, the total air space sampled by the Townes trap

was about three-fourths that of the Cornell trap. Once each week,

the positions of the two traps in each location were switched to

control for trap placement effects. All traps used were in their

second or third season of operation.

IManufactured by Cornell Eavipment Co. Ine., 1115 BW. Rolling. Rd...

Baltimore, Maryland 21228

430 Contribs” Amer.«Ent.: Insts, vob. 20, 1983

RESULTS

The 4 traps collected 69,247 insects over the 4 weeks of the

experiment (Table 1), the Townes traps accounting for just over 90%

of this total. The Townes traps caught over ten times as many

Diptera, almost six times as many Hemiptera, and three and one-half

times as many Lepidoptera as the Cornell traps. Similarly, the

Townes traps caught 88.7% of all Hymenoptera obtained during the

study. The two trap types caught roughly equivalent percentages of

the various hymenopteran taxa, however. Ichneumonidae accounted

for approximately 61% and Braconidae, approximately 28%.

Although traps were placed side by side, the alternate

positions at each site were not equivalent (Table 2). For three of

the four comparison pairs, the differences in numbers of insects

caught were highly significant; only the Cornell trap at the woods

site caught equal numbers of insects in the two alternate

positions.

TABLE 1. TOTAL CATCHES OF COMMERCIAL CORNELL AND TOWNES MALAISE TRAPS

COMPARED FOR PAIRS OF TRAPS AT TWO LOCATIONS FOR FOUR WEEKS. FOR

ADDITIONAL DETAILS SEE TEXT.

Taxon Trout Pond Ash Woods Overall

Commercial __Townes Commercial _Townes Totals

Diptera 4, 204( 74.4%) 34,425(83.2%) 712(56.9%) 16,265(77.6%) 55,606

Lepidoptera 480 (8.5%) 1,437 (3.5%) 149(11.9%) Fa6.-63.5%) - 92,792

Hymenoptera 361 (6.4%) 1,792 (4.3%) 270(21.6%) 3,137(15.0%) 5,560

Other orders 172 (3.0%) 1,345 (3.3%) 86 (6.9%) 485 (2.3%) 2,098

Total

Insecta 5,652 41,394 1,254 20,960 69,247

TABLE 2. TOTAL CATCH IN RELATION TO TRAP POSITION AT EACH OF THE

TWO STUDY SITES ACCORDING TO THE KIND OF TRAP USED.

Trap 7 Trout Pond Chi square Ash Woods Chi square

Type Pos. A Pos. B Value Pos, A Pos, B NMalue

Commercial 3,337 2,315 184.8* 595 656 3.0

Townes 2b 867, 16, 5e%o 19676 07? 10,201 «105759 10:3 9°

Combined

Traps 26,794 ' 167,682” 485900" 10,796 11,415 4703"

Co ee ee ae

Significantly different, p<.001.

Matthews and Matthews: Malaise Trap 431

DISCUSSION

Two facts emerge from this study. First, regardless of trap

position or placement, the Townes traps were vastly superior to the

commercial model by every measure of collecting efficiency.

Second, even a relatively small change in trap placement resulted

in large differences in collection efficiency.

It is well known that insect flight tends to be concentrated

along certain natural corridors such as streams, ecotones, and

woodland paths. For Malaise traps to be maximally effective, they

should be located in such flyways. The narrow nature of these

flight corridors is well illustrated by our data. Although at each

site we attempted to place both traps within a single flyway, the

two side-by-side locations still differed significantly in numbers

of insects obtained. These data underscore the caution with which

researchers should attempt any use of Malaise traps for comparing

different habitats.

The variables that make one trap design more efficient than

another are as yet poorly understood. Because of differences in

the size of the trap openings, the commercial trap would

superficially appear to sample a larger air space (4m2) than the

Townes trap(3m@). However, the four-sided design of the commercial

trap probably makes such a comparison misleading. When such

Malaise traps are placed ina flyway, the two sides parallel to the

insect flight path are probably nearly functionless. Thus, the

effective air space of the trap is more likely to be on the order

of 2m? than 4m2@, However, even when our commercial trap data are

corrected for this, the Townes traps remain superior in numbers of

insects caught.

Trap color is known to affect catch (Roberts, 1970,1975;

Townes, 1972). The bicolor design of the Townes trap probably

increased its effectiveness. Green traps have been shown to trap

fewer tabanid flies than natural saran (Roberts, 1970). Further

Studies on this aspect of trap design might be enlightening.

In our comparison, the shape of the trap openings which

intercept insect flight may also have been a major contributing

factor to the differences obtained. In the Townes model, these

openings are roughly rectangular, whereas in the commercial trap

they are triangular. Thus, while both traps intercept much the

Same area at ground level, they differ more and more at higher

levels. (Thus, one would expect a more detailed analysis of our

comparative data to reveal that the commercial trap weighted

Samples in favor of those species that tend to fly close to the

ground. )

The bias inherent in the triangular trap opening of the

commercial trap probably also Fras seasonal implications. If height

of insect flight above the ground varies according to seasonal

temperature (Townes, 1962), the differences in trap efficiency

would be accentuated during the warmest periods when insect flight

traffic is concentrated higher above the soil. support for this

idea comes from our use of these commercial traps continuously over

13 weeks in Summer 1967 (see Fig. 2, Matthews and Matthews, 1970)

in the same positions and locations as those reported here; the

greatest numbers of insects were captured during the early weeks of

Summer, declining as the climate warmed.

432 Contrib. Amer. Ent. Insts, vol. .20, 1983

ACKNOWLEDGEMENTS

This research was supported by the E. N. Huyck Preserve,

Rensselaerville, New York. We extend our grateful appreciation to

the Preserve staff, and especially to Dr. Robert Dalgleish and

Sally Hoyt. Special thanks to Wayne Gardner, Kathryn Jablonski,

Ken Suarez, Kent Palmer, and Mikko Bojarsky, enthusiastic members

of two EARTHWATCH teams who assisted in this study.

LITERATURE CITED

Gressitt, J. L. and M. K. Gressitt. 1962. An improved Malaise

trap. Pacific Insects 4: 87-90.

Matthews, R. W. and J. R. Matthews. 1970. Malaise trap studies of

flying insects ina New York mesic forest. I. Ordinal

composition and seasonal abundance. Veet. ait Soo. Aebs

52-59.

1971. The Malaise trap: Its utility and potential

for sampling insect populations. Mich. Ent. 4: 117-122.

Roberts, R. H. 1970. Color of Malaise trap and the collection of

Tabanidae. Mosq. News 30: 567-571.

1975. Influence of trap screen age on collections

of tabanids in Malaise traps. Mosq. News 35: 538-539.

Steyskal, G. C. 1981. A bibliography of the malaise trap. Proc.

Ent. Soc. Wash. 83: 225-229.

Townes, H. 1962. Design for a Malaise trap. Proc. Ent. Soc.

Wash. 64: 253-262.

1972. A light-weight Malaise trap. Ent. News 83:

239-247.

BIOLOGICAL STUDIES ON GonrIOZUS LEGNERI GORDH (HYMENOPTERA:

BETHYLIDAE) A PRIMARY EXTERNAL PARASITE OF THE NAVEL

ORANGEWORM AMYELOIS TRANSITELLA AND PINK BOLLWORM

PECTINOPHORA GOSSYPIELLA (LEPIDOPTERA: PYRALIDAE, GELECHIIDAE)

Gordon Gordh, James B. Woolley and R. A. Medved

Division of Biological Control, Department of Entomology

University of California, Riverside, California 92521

Abstract

Goniozus legneri Gordh is a primary external parasite of Microlepidop-

tera larvae (Gelechiidae, Pyralidae), which was imported into California from

southern Uruguay for the biological control of Navel Orangeworm (NOW),

Amyelois transitella (Walker), a serious pest on almonds. Details of the

biology of this parasite are described and compared with its performance on

the Pink Bollworm (PBW), Pectinophora gossypiella (Saunders). Studies

showed that on NOW, G. legneri attacked 17.23 + 2.97 hosts to sperm deple-

tion, attacked 6.12 + 3.34 hosts after sperm depletion, and parasitized 23.35

+ 2.58 hosts at-25 (+..2°C;.50. 22%. RH, LD.44210-(m =.26). On PBW, Gs.)

legneri attacked 21.45 + 3.91 hosts to sperm depletion, and 25.05 + 3.55 dur-

ing the life of the female parasite under the same environmental conditions

(n = 20). Differences between means were not statistically significant.

On NOW, sting to oviposition time was 1.19 + 0.47 days for broods pro-

ducing male and female progeny (n = 437, and 1.82 + 1.16 days for broods

producing only male progeny (n = 159). The period from oviposition to eclo-

sion was 1.25 + 0.27 days for broods producing male and female progeny

(n = 436), and 1.23 + 0.27 days for broods producing only males (n = 155).

Development time from larva to cocoon was 3.75 + 0.35 days for broods of

males and females (n = 430), and 3.60 + 0.21 days for broods of only males

(n = 155). Development time from cocoon to adult was 9.26 + 0.53 days for

broods producing male and female progeny (n = 430), and 8.63 + 0.40 days

for broods producing only male progeny (n = 155). Mean development time

from egg to adult for broods of male and female parasites was 14.26 + 0.97

rod (n y 430), and 13.45 + 0.46 days for broods composed of only males

n° * 195).

On PBW, sting to oviposition time was 1.40 + 0.59 days for broods pro-

ducing only male and female progeny (n = 429), and 1.96 + 1.07 days for

_ broods producing only male progeny (n = 71). The period from oviposition

to eclosion was 1.44 + 0.18 days for broods producing male and female

progeny (n = 428), and 1.46 + 0.13 days for broods producing only male prog-

eny (n = 71). Development time from larva to cocoon was 3.61 + 0.26 days

for broods producing male and female progeny (n = 428), and 3.58 + 0.23

days for broods producing only male progeny (n = 71). Development time

from cocoon to adult was 8.93 + 0.57 days for broods producing male and

female progeny (n = 428), and 8.96 + 0.38 days for broods producing only

male progeny (n = 71). Mean development time from egg to adult was

13.98 + 0.53 days for broods producing male and female progeny (n = 427),

and 14.00 + 0.47 days for broods producing only male progeny (n = 7).

The egg and larval stages are described; they do not differ from other

species of Goniozus which have been examined. The pattern and sequence of

433

434 Contrib. Amer. Ent. Inst., vol. 20, 1983

pupal pigmentation is described. This character may be important in recog-

nizing subtle morphological differences among closely related species or pos-

sibly has phylogenetic significance.

Goniozus legneri prefers to oviposit on the dorsal aspect of the host's

body, followed by the lateral and ventral surfaces. On NOW, segment pref-~

erence is 5-9 for all aspects, with a normal distribution peaking on seg-

ments 7-8 for all aspects. Segment and aspect preference is identical on

PBW. This data was compared with data collected in a similar manner for

G. gordhi Evans and G. aethops Evans.

Goniozus legneri has a precise sex ratio. The sex ratio tends to re-

main constant over a wide range of host weights with supplemental males

produced on heavier hosts which, presumably, can sustain larger broods of

parasite progeny without superparasitism.

We found that on NOW (n = 26) and PBW (n = 20) G. legneri has a

normal distribution of eggs on the host body and that when host weight

varies between 20.0 and 50.0 mg, on PBW there was a steady increase in the

number of eggs laid per host with increasing host weight with a maximum of

16.0 eggs per host in the largest host weight class observed (42.5 - 45.0 mg).

On NOW oviposition increased with increasing host weight to a maximum of

18.0 eggs per host in the 45.0-47.5 mg host weight class for female para-

sites provided with a 24 hour interval between ovipositions, and reached a

maximum of 13.2 eggs per host in the 37.5 -40.0 mg host weight class and

then remained relatively constant (ll.1-13.0 eggs per host) for females pro-

vided with fresh hosts immediately following oviposition. Fewer eggs laid on

lighter hosts suggest that the parasite can regulate the number of eggs laid

to prevent superparasitism; a relatively constant number of eggs laid at higher

host weights suggests a depletion of fully developed eggs. These data were

compared with other species of Goniozus. |

Life tables were constructed for G. legneri on NOW and PBW, an

demographic statistics were computed and compared with those generated

from a life table analysis of G. emigratus reared on NOW. Intrinsic rates

of increase Lf for G. legneri on PBW and NOW, and G. emigratus on

NOW were 0.157, 0.162, and 0.178 females per female per day, respectively.

Capacities for increase (r¢) were substantially less than corresponding r

values: 0.0985 and 0.110 females per female per day for G. legneri on

PBW and NOW, respectively; generation times (T,) were 51.5 and 45.1 days,

respectively; and net reproductive rates (Rg) were 160.2 and 145.1 females

per female per lifetime, respectively.

Longevity studies showed that mated females deprived of hosts lived

75.0 + 5.81 days (n = 50) at 25 + 2°C, unmated females lived 71.0 + 10.8]

days (n = 50), and unmated males lived 61.3 + 15.87 days (n = 30) under

the same conditions. Longevity of mated female parasites on PBW was 82.35

+ 14.74 days. Regarding host survival time, one set of 10 NOW hosts was

provided female G. legneri which stung the host and oviposited on the NOW

larvae. Eggs hatched and larval development was completed. Based on

haemolymph flow observed and dorsal aorta contraction, longevity of the host

was 5.0 + 0.00 days at 25 + 2°C, 50 + 2% RH, and LD 14:10 (n = 10).

Pulse rates of parasitized NOW were compared during the morning and even-

ing, these were not significantly different for each day of life but were

significantly different on the first and last day of host life. Similar

Gordh, Woolley & Medved: Biology of Goniozus 435

information was collected for paralyzed, but not parasitized hosts. Paralyzed

hosts lived 29.20 + 6.49 days (n = 10) under the same environmental condi-

tions. Data on pulse rate (morning vs afternoon, first day after sting and

last day of life) is presented.

A stepwide regression procedure was used to analyze sex ratio of the

first 12 broods of G. legneri attacking NOW (n = 6) and was performed for

sex ratio (dependent variable) against six independent variables (weight of the

host, age of the parasite, number of eggs deposited, days since the last

sting, female adult longevity, and brood mortality). The regression was

statistically significant for sex ratio and days to last sting (i.e., days since

the last host was attacked) with an r2 of 0.0811 and an F statistic of 6.18

(p = 0.0153, 70 degrees of freedom).

Similarly, a correlation analysis was conducted on 26 female G.

legneri attacking NOW and 20 females attacking PBW. Correlation coeffi-

cients for egg, larval and pupal mortality on a given host were not statistic-

ally significant. :

INTRODUCTION

This report concerns the biology of Goniozus legneri Gordh, a member

of the Bethylidae. Bethylids presumably represent a primitive family of

aculeate Hymenoptera. The family now consists of about 40 valid genera

and about 2,000 nominal species. The Nearctic species have recently been

revised by Evans (1978b), and he has also treated the Neotropical species

(1962; 1964; 1969 a,b,c; 1970; 1973; 1978a; 1979 a,b). Substantial contributions

on the Palearctic and Ethiopian faunas have been made by Moéoczar (1966;

1969; 1970 a,b,c; 1971; 1974; 1975; 1976 a,b; 1978; 1981) and Nagy (1966, 1967;

1968 a,b,c; 1969; 1970 a,b; 1972; 1974; 1976). A world catalog is in prepara-

tion (Gordh and Hawkins).

All bethylids are primary external parasites of Microlepidoptera and

Coleoptera larvae. There are no known bethylid hyperparasites. Due to their

host associations they are of potential benefit in the biological control of

agricultural and stored product pests (Chatterjee 1941, Cushman and Gordh

1976, Gordh 1976, Gordh and Evans 1976, Gordh and Hartman in press, Gordh

and Hawkins 1981, Kishitani 1961, Kuhne and Becker 1974, Mertens 1981,

Remadevi et al. 1978, Venkatraman and Chacko 1961, Voukassovitch 1924,

Yamada 1955).

The genus Goniozus is probably the most well studied genus of

bethylids and displays excellent promise for biological control of Microlepidop -

tera. The genus is cosmopolitan in distribution and contains more than 50

nominal species. Of the species which have been studied, it appears partic-

ularly effective on Gelechiidae and Pyralidae. No species has been demon-

strated to be host specific.

Goniozus legneri was imported by Dr. E. F. Legner into California

from southern Uruguay for the biological control of Navel Orangeworm,

Amyelois transitella (Walker) (hereafter designated NOW). This pyralid

is the most serious orchard pest of pecans and almonds in the Central

Valley of California. Initial importation was made in 1978, and releases were

made in the Central Valley in 1979. The parasite was recovered in the

436 Contrib. Amer. Ent. Inst., vol. 20, 1983

spring of 1980 and innoculative releases were made in 198] and 1982. Studies

by Legner (Legner et al. 1982 a,b) have shown that the parasite is estab-

lished and that it is an important component in the natural enemy complex

of NOW on almonds on study sites in the Central Valley. This paper pro-

vides laboratory-generated biological information on the parasite which sup-

plements field-collected data by Legner, and provides new information

intended to supplement existing knowledge on the genus Goniozus.

Voucher specimens of G. legneri have been deposited in the following

institutions: Beneficial Insects Introduction Institute, Beltsville, MD; Bio-

systematics Research Institute, Agriculture Canada, Ottawa; Zoological Insti-

tute, Academy of Sciences of the U.S.S.R., Leningrad; California Academy of

Science, San Francisco; All Union Plant Protection Institute, Pretoria, South

Africa; Faculty of Agriculture, Ehime University, Shikoku, Japan; Department

of Entomology, University of Florida, Gainesville; Department of Zoology and

Entomology, Fort Collins, Colorado; University of California, Riverside, Ento-

mology Department; Queensland Museum, Brisbane, Australia; American Mus-

eum of Natural History, New York.

MATERIALS AND METHODS

Goniozus legneri was reared in the laboratory on NOW and Pink Boll-

worm, Pectinophora gossypiella (Walker) (hereafter designated PBW).

Individual parasites were placed in four-dram, plastic snap-cap vials with a

few droplets of honey provided as a supplemental source of nutrition. Vials

were lodged in a Percival Temperature Cabinet set at 25 + 2°C, 50 + 2% RH,

and LD 14:10. Experience has shown that containers must have extremely

tight fitting lids because the parasites are fossorial and will escape from

other types of containers.

To determine parasite longevity, we performed the following experiment.

Broods which developed on NOW were followed through to adult emergence

and individuals placed in four-dram, snap-cap vials with a few droplets of

honey provided as a supplemental source of nutrition. Two experimental

groups were used, with fifty replicates per group. One group was provided

with hosts (NOW and PBW, 50 replicates each) and one group was deprived

of hosts. The group deprived of hosts consisted of 50 mated and 50 unmated

female parasites. Additionally, we included a group of 30 unmated male

G. legneri in the experiment. The vials were placed in a Percival Cabinet

set at environmental conditions specified earlier. On all experimental

studies, parasites were newly emerged and copulation had been observed.

In subsequent studies we were interested in examining several aspects

of the biology of G. legneri. In the longevity study we learned that 50

replicates per experiment were unmanageable from the standpoint of main-

taining host cultures and studying parasite biology (time required to collect

data). Thus we conducted a detailed analysis of the biology of G. legneri

using 26 females on NOW and 21 females on PBW. Initially, 30 parasites

were set up on NOW, but four females were killed or escaped during host

transfer. We decided to set up 21 parasites on PBW because of problems

with the host culture. PBW is cannibalistic under crowded conditions and it

was not always possible to maintain the culture at high density, or a

Gordh, Woolley & Medved: Biology of Goniozus 437

population density necessary to conduct experiments under conditions similar

to those used for NOW.

Individual female parasites were isolated in numbered vials in the above

mentioned manner and given one fourth instar host at a time. The hosts

were weighed with a Metler analytical balance to the nearest tenth of a mil-

ligram before being offered to female parasites. The following information

was tabulated for each parasite: (1) time of paralysis (day since emergence

or day since last sting); (2) time interval between paralysis and oviposition;

(3) number of eggs laid and the position of eggs on each body segment of

the host (i.e., dorsal, lateral, ventral); (4) orientation of the paralyzed larva

in the container (i.e., dorsal, lateral, ventral); and (5) extent of host feeding

by the adult female parasite. In no instance was an alternate host species

provided to a female parasite attacking a given host (NOW or PBW).

Parasitized hosts were isolated in 000 size gelatin capsules (Lilly Phar-

maceutical Company) and the capsules were sequentially taped on 5 x 8 inch

cards and placed in an environmental control cabinet under the conditions

specified above. Daily observations were made of this material and progeny

development time, sex ratio of progeny, and immature mortality were tab-

ulated and computed. In turn, this information was correlated to adult para-

site age and the number of previous reproductive episodes for each female of

the parental generation. After each reproductive episode the adult female

parasite was isolated for 24 hours (deprived of hosts, but not honey) and

then given another 4th instar host. This procedure was repeated throughout

the lifetime of each parasite on either host.

Following the deaths of the 26 females on NOW and the 2] females on

PBW, life table statistics were calculated for G. legneri on each host using

the computational methods of Andrewartha and Birch (1954) and Birch (1948).

Age was measured in days. The counts of live Ist instar parasite larvae 24

hours after oviposition were used to estimate egg viability. The number of

viable female eggs was counted (in egg clutches with no further mortality)

or estimated (in clutches with larval or pupal mortality) from the number of

adult female progeny in each egg clutch. Age specific fecundity (m,) was

calculated as the number of viable female eggs produced per day divided by

the number of females alive on that day. Daily survivorship (1,) for the

immature stages of the parental females was estimated from that noted in

their progeny. Daily survivorship (1,) for adult females was calculated

directly as the number of females surviving over the initial number of females

times the estimated survivorship at adult emergence.

Hosts were cultured in the following manner. The NOW culture was

about 2.5 years and came from field-collected material taken in the Central

Valley. Adult moths were placed in 0.5 gallon ice cream containers fitted

with wire screen lids. Filter paper was placed on the screens and the

moths oviposited on the paper. Eggs were treated in 10% formalin and then

rinsed in cool running water for 45 minutes. The sheets were then hung to

air dry. First instar larvae were transferred with a 0000 size camel's hair

brush to one ounce jelly cups filled with artificial media. The media con-

sisted on two parts honey, two parts glycerine and one part water (total

volume 250 ml). Two teaspoons of yeast and six grams of vitamins were

then mixed into the solution and then poured over 1200 ml of bran. Develop-

ment time of the hosts required about 28 days at 25°C.

438 Contrib. Amer. Ent. Inst., vol. 20, 1983

The PBW culture was about one year old and also was from field-

recovered material. Females were placed in 0.5 gallon ice cream containers

fitted with wire screen lids and filter paper used as an ovipositional sub-

strate. After egg hatch the first instar larvae were transferred to one

ounce jelly cups filled with an artificial media. The media consisted of 75

gr Agar, 180 gr Casein, 50 gr salt, 180 gr sugar, 25 gr Alphacel, 8 gr MPH,

150 gr wheat germ, II5 gr vitamins, 15 ml 40% formaldehyde, 25 ml 10% KOH,

and 55 ml acetic acid. The mixture was blended and boiled and put in the

jelly cups and allowed to cool and solidify before the larvae were transferred.

About 21 days at 25°C were necessary for pupation.

RESULTS

1. Parasite Longevity: Results of the parasite longevity experiment are

indicated in Figure 1. Mated females deprived of hosts lived 75.0 + 5.8l

days; unmated females deprived of hosts lived 71.0 + 10.81 days. The differ-

ences were statistically different at P = 0.05. We attributed this difference

to unmated females wandering around the container and in general being

more active. Unmated males lived 61.3 + 15.87 days, and this was also sta-

tistically significantly different from mated and unmated females at P = 0.05.

Mated female parasites provided PBW hosts throughout their lifetime

lived 95.76 + 10.53 days. Mated female parasites provided NOW lived 97.35

+ 14.74 days. These results were not statistically significantly different, but

were significantly different from the host deprived group (P = 0.01). We sus-

pect that host feeding and/or reproductive activity may be involved in the

observed differences. When daily observations were made, it appeared as if

the host deprived class was visibly more active. This aspect of parasite

longevity will be investigated in a subsequent study.

2. Parasite Host-attack: We found that on NOW the 26 parasites

attacked 23.35 + 2.58 hosts each, and that on PBW the 21 parasites attacked

24.76 + 3.81 hosts each. The differences between hosts attacked were not

statistically significant. Based on the sex ratio of progeny produced, we

found that G. legneri attacked 17.23 + 2.97 hosts until sperm depletion

(i.e., on the average 17 broods produced females and that subsequent hosts

produced all male broods) on NOW and on PBW the females attacked 21.38

+ 3.82 hosts until sperm depletion. The differences are not statistically dif-

ferent, but the slightly higher number noted for PBW was attributed to the

fact that it is a slightly larger host and a few more males were produced

per brood; the parental stock used in the study was generated from sib

mating. That is, replicates represented all of the females mated to

brothers issuing from a few clutches of eggs. From these data we noted

that on NOW mean female progeny production was 149.2] + 17.62 per paren-

tal life time while parental females attacking PBW produced 169.55 + 28.80

female progeny. Again, we interpret the higher production of females to

the fact that more males per brood were produced on PBW, hence, on PBW

an individual male had more sperm available for each ejaculation. Goniozus

legneri is arrhenotokous; unfertilized eggs produce males, fertilized eggs

produce females. Thus it would appear that G. legneri averages at least

150 sperm on NOW and about 170 sperm on PBW. The rather high standard

deviations may be attributable to variation in ejaculate size in the mating

Gordh, Woolley & Medved: Biology of Goniozus 439

sequence (sister number one getting more sperm than sister number six),

size of the spermatheca limiting sperm accommodation, or both.

3. Host Attack and Oviposition: Immediately after adult emergence,

G. legneri has a preovipositional period of 3.12 + 0.67 days on NOW (n =

26), and 4.05 + 2.16 days on PBW (n = 20). During this period the parasite

is capable of stinging and paralyzing the host larva, but it is not capable of

oviposition. Dissection of newly emerged female parasites shows that the

ovaries are developed, but that eggs are not.

We have not determined the behavioral modalities used in host finding.

Under the experimental conditions used, the parasite seemed to wander ran-

domly in the arena until the host was located. Once detected, the para-

site quickly mounted the host. The attack is rather specific. The female

attaches to the host's integument with her mandibles embedded in the dorsum

just posteriad of the head capsule. She then thrusts her metasoma ventrad

and stings the host, presumably in the ganglion of the ventral nerve cord just

posteriad of the gula. We believe the site is important because in many

instances the parasite would repeatedly thrust her sting while the host larva

writhed in an attempt to dislodge the parasite. In other instances one thrust

was sufficient to immediately paralyze the host. In some instances total

paralysis required about one minute during which the female parasite waited

3-4 cm from the host, grooming herself. We found that 90 day old,

senescent parasites incapable of oviposition were capable of paralyzing their

NOW hosts. Although the paralysis prevents locomotion, the paralyzed host

is still capable of lateral flexion when stimulated with a probe. Physiolog-

ically, the venom is interesting because if the host was paralyzed just prior

to the time of pupation, the host pupated and eggs deposited on such hosts

did not develop.

We analyzed host survival based on the number of contractions per

minutes of the dorsal aorta of paralyzed versus paralyzed and parasitized

hosts (Fig. 2). Two groups of ten newly emerged G. legneri were allowed

to paralyze NOW larvae and one group was allowed to parasitize their hosts.

The second group was removed after host paralysis, but before oviposition.

The pulse rate was counted each morning and evening for each paralyzed and

paralyzed and parasitized host. There was no significant difference between

morning and evening counts, but within the paralyzed and parasitized group

there was a linear decline in pulse rate over five days, with all hosts "dead"

(no haemolymph flow) on the sixth day. The variance in haemolymph flow

among hosts in this group was nearly constant (i.e., not significantly differ-

_-ent among individuals).

Hosts which were paralyzed and not parasitized lived 28.30 + 6.65 days,

based on contraction of the dorsal aorta. After an initial decline and up-

shift of heart beat rate, pulse rate remained relatively constant between the

10th and 30th days. The variance was rather constant until the 24th day,

after which it became somewhat erratic. The temperature was constant

(25°C) throughout the experiment, so the increase in variance was attributed

to a smaller sample size and varying degrees of moribundity within the

sample population.

_Oviposition requires two to four hours. The period from sting to ovi-

position requires 1.19 + 0.47 days for females producing male and female

440 Contrib. Amer. Ent. Inst., vol. 20, 1983

progeny, and 1.82 + 1.16 days for females producing only male progeny on

NOW (n = 437, 159, respectively). On PBW, sting to oviposition was 1.40 +

0.59 days for females producing male and female progeny, and 1.96 + 1.07

days for females producing only male progeny (n = 429, 71, respectively).

The differences between hosts were not statistically significant, but females

producing only male progeny were old (ca. 50-60 days) and had depleted

their sperm supply. Presumably their senescent condition was responsible for

the longer sting-to-oviposition period. Summary data on oviposition to eclo-

sion, larval development to cocoon formation, cocoon to adult emergence

(pupation), and oviposition to adult emergence is provided for parasites at-

tacking NOW and PBW (Table 1).

Occasionally, a female may leave a host for a few minutes or become

quiescent, but she returns to the host or becomes ovipositionally active until

the appropriate number of eggs have been deposited on the host (see below).

Once the full complement of eggs has been deposited, the female leaves the

host and wanders around the arena. If an ovipositing female is confronted

with a pencil or probe, she will act aggressively toward it by spreading her

mandibles and attempting to bite or sting the object. Similar behavior has

been observed in females near paralyzed and arasitized hosts.

Eggs may be deposited one hour after paralysis of the host or the

parasite may wait up to five days. The period from oviposition to eclosion

was 1.25 + 0.27 days for broods producing male and female progeny on NOW

(n = 436) and 1.23 + 0.27 days for broods producing only male progeny

(n = 155). Parasite eggs on PBW required 1.44 + C.18 days to develop for

broods producing male and female progeny (n = 428), and 1.46 + 0.13 days

for broods producing only males (n = 71). We are unable to explain the

slightly longer period of the egg stage on PBW. In all instances observa-

tions were made in the morning and evening.

During the period of oviposition the host remains alive and haemolymph

flows metasynchronously in the dorsal aorta. Females will host feed before

oviposition and frequently necrotic areas on the host's integument appear at

the site of feeding. Males have not been observed to host feed.

The larval to cocoon interval was slightly shorter for all male broods,

for broods on NOW (3.75 + 0.35 (n = 430) versus 3.60 + 0.21 (n = 155) and

PBW (3.61 + 0.26 [n = 428] versus 3.58 + 0.23 [n = 71]), but the differences

were not statistically significant. Males are invariably smaller than females

and emerge before sibling sisters. It seems that larval stage is responsible

for this difference between sexes, although it is not statistically significant.

Males emerge less than 12 hours before females of the same brood, but the

data was collected twice daily. This could have a confounding effect on the

Statistics presented in Table 2, as more frequent observations might have pro-

duced a statistically significant results.

The larval head of G. legneri does not project through the hosts's

integument but rather only the mandibles are embedded in the cuticle. As

with the egg, removing the parasite larva from the host results in its death.

During the first two days of larval development the parasites consume about

50% of the internal contents of the host. Development time for the larval

stage for mixed and male only broods on both hosts at 25°C and 50% RH are

given above and in Table 1. The larvae eventually consume the contents of

Gordh, Woolley & Medved: Biology of Goniozus 441

the host's body and all that remains is a shriveled cuticle and the head cap-

sule. We believe that haemolymph flow is essential to parasite development

because in several instances when the adult female parasite killed the host

(determined by lack of contraction of the dorsal aorta) and oviposited upon

it, the eggs would hatch but invariably the larvae died.

After completing feeding, the larvae detach from the host but remain

in situ and begin to spin a cocoon. The spinning larva differs from the

feeding larva in that the integument is no longer transparent and urate cells

aggregate near the integument and form white spherules.

The cocoon of G. legneri is loosely woven, white, 5 mm long and 1.5

mm in diameter. Its construction requires 14-16 hours at 25°C. After com-

pletion of the cocoon a dark orange fluid is secreted from the anus. The

amount of time between the onset of spinning and voiding the fluid is about

30 hours. The fluid hardens in several minutes, depending on ambient temper-

ature and relative humidity. Stimulation, such as probing or squeezing with

a pair of forceps causes the hindgut to be voided. In all instances, however,

the hindgut is voided in less than five seconds.

The pupal period, defined as the termination of cocoon formation to

adult emergence, is 9.26 + 0.53 days on NOW (n = 430) at 25°C and 50%

RH, for female and male broods, and 8.63 + 0.40 days for broods producing

only male progeny (n = 155). On PBW pupal development requires 8.93 +

0.57 days for broods producing males and females (n = 428), and 8.96 + 0.38

days for broods composed exclusively of males (n = 71). The differences

between PBW and NOW were different, but not statistically significant.

However, the difference between the means for mixed and all male broods

on NOW is highly significant (p < 0.001). This was to be expected because

males are protandrous and emerge before their sibling sisters. Mixed broods

would be expected to have a longer mean development time. There was no

significant difference in the period of pupation when progeny from a young

parasite was compared to the progeny of an old parasite on either host. In

fact, we found no statistically significant correlation between age of the

parental female and progeny development.

The newly formed pupa is entirely white with translucent appendages.

Pigmentation occurs in a definite, predictable pattern (Fig. 3). Initially the

eyes become pink. As they darken the dorsum of the propodeum and scutel-

lum also darken. Ventrad, the middle and hind coxae darken and the medial

portion of the sternum becomes dusky. Later the darkening of these areas

becomes more pronounced and the posterior portions of metasomal sterna I-

III, and the ventral aspect of the head becomes dusky. Darkening of the

above mentioned regions proceeds and spreads such that within 96 hours the

entire body assumes adult coloration.

Total development time from egg to adult was 14.26 + 0.97 days on

NOW at 25°C and 50% RH for broods producing males and females (n = 430).

Development time under the same conditions for broods producing only

males was 13.45 + 0.46 days (n = 155). The difference is highly significant

(p < 0.001) and due to the combined effect of shorter larval and pupal

periods for all male broods. Development time for broods producing males

and females on PBW was 13.98 + 0.53 days (n = 427), and 14.00 + 0.47 days

for broods producing only males (n = 71). The differences between

442 Contrib. Amer. Ent. Inst., vol. 20, 1983

development times for equivalent broods on NOW versus PBW are highly

significant (p < 0.001) but opposite in sign and are attributable to the larval

and pupal development periods. We have no explanation for the high vari-

ance in development time (oviposition to adult) for mixed broods developing

on NOW.

4. Courtship, Copulation, and Sex Ratio: Males of G. legneri are

protandrous and emerge 10-12 hours before their sibling sisters. After emer-

gence, a male remains at the site of pupation and chews a hole in the fe-

male's cocoon. No particular site is preferred for the hole and the male

may chew into the anterior or posterior end of the female's cocoon. Once

inside the cocoon, the male orients toward the female head-to-head and

dorsum -to-venter, with the male assuming the superior position. The female

will deflect her metasoma in an upward position and the male will extend

his metasoma to the ventral surface of the female's metasoma. Little

overt courtship behavior exists in this species and copulation requires 3 -15

minutes. The seemingly wide variation in copulation time is probably due to

female receptivity, male sperm depletion (ejaculate size), or both. After

inseminating the female, the male leaves the cocoon and chews a hole in

another female's cocoon and the procedure is repeated. In the meantime,

the inseminated female leaves her cocoon and grooms her body. Copulation

also occurs between males and newly emerged, uninseminated females.

Inseminated females repel subsequent copulatory attempts by males.

If a male succeeds in mounting an unreceptive female, she runs rapidly

around the container. If this behavior does not repel the male, she uses

her hind legs in an attempt to dislodge the male. Frequently, females

will turn in an attempt to bite the male.

In one experiment, cocoons of both sexes were isolated and placed in-

dividually in 000 gelatin capsules. After ten days individual virgin males and

females were placed together in four-dram vials. Observing ten such couples

revealed that not all females were readily receptive to courtship and copu-

latory attempts of virgin males. Receptive females remained quiescent with

their antennae vibrating. Insemination in these episodes lasted three to five

minutes.

Goniozus legneri has a precise sex ratio and regulates the number of

male eggs based on the size of its host. The overall sex ratio for 26 fe-

male parasites throughout their lifetime on NOW (including clutches which

produced only males) was 1.23:1.00 in favor of females (n = 6,761 eggs).

However, when only the first twelve broods are considered, i.e., ignoring

all-male broods produced by females which have depleted their sperm supply,

the sex ratio shifts to 4.15:1.00 in favor of females.

In order to analyze the effects of various factors on sex ratio a step-

wise regression procedure was performed on the data for the first 12 broods

of G. legneri on NOW (n = 6). Sex ratio (as number of females over the

total brood, arcsin transformation) was treated as the dependent variable

against six independent variables: host weight, age of the parasite, number

of eggs in the brood, days since the last sting, female adult longevity, and

brood mortality. The regression was statistically significant for sex ratio and

days since last sting with an r2 of 0.081 and an F statistic of 6.18

Gordh, Woolley & Medved: Biology of Goniozus 443

(p = 0.0153, 70 degrees of freedom). That is, the proportion of females in

a brood tends to increase as the number of days since the last oviposition

increases. A longer period between oviposition may provide for increased

sperm availability during the next reproductive episode.

Table 2 represents a joint frequency distribution for the first 12 broods

of 191 clutches of eggs of G. legneri laid on NOW for which there was no

immature mortality. From the data presented it is obvious that the number

of males produced is strongly correlated with brood size. Small broods have

only one male; intermediate sized broods have two males while large broods

produce three males. It seems apparent that supplemental males are pro-

duced for intermediate and large broods to insure adequate sperm to insemi-

nate all females of the brood. We have observed numerous inseminative

episodes in multiple male broods and have not witnessed combat among males

for uninseminated females in a brood.

5. Egg Production and Host Size: Studies with other species of

Goniozus suggest that the number of eggs per clutch is strongly correlated

with host size. In order to evaluate the relationship between host size and

clutch size we used the data from the detailed tests of 26 G. legneri

females on NOW and 20 G. legneri females on PBW (the 2lst female on

G. legneri was originally set up as an alternate in case of parasite mortal-

ity. Thus, the data from this female was not included in some analyses.

It was in no way atypical). These parasites were provided with weighed

hosts following a 24 hour isolation interval after each oviposition, as de-

scribed above. Hosts were chosen at random from host cultures. In addi-

tion, concurrently with the 26 females on NOW described above, 6 females

were set up on NOW under identical conditions with the exception that

these 6 females were given fresh hosts immediately after oviposition was

noted (i.e., no 24 hour isolation interval). Due to logistical reasons, the

NOW and PBW tests could not be run concurrently. However, the repli-

cates were exposed to identical conditions in all tests.

The weight of hosts was broken down into 2.5 mg increments and the

total number of parasite eggs, the mean number of eggs per host (clutch),

and the number of such clutches observed was tabulated for each host

weight class for each of the 3 tests. The data are presented in Table 3.

The distribution of host weights for NOW and PBW is apparently nor-

mal, as one would expect, with the largest number of hosts in the ranges

22.5 to 35.0 mg for PBW and 25.0 to 40.0 mg for NOW. The absolute num-

ber of eggs in each weight class follows these distributions. However, the

mean number of eggs per host shows an increase with increasing host weight

for PBW and for NOW with and without a 24 hour isolation interval. The

trend is monotonic with one exception on PBW, and on this host the number

of eggs per host reaches a maximum in the largest weight class provided.

It is unknown, of course, if clutch size on PBW would increase if larger PBW

larvae were available. On NOW, however, the data for clutch size suggest a

plateau in the weight classes of 45.0 to over 50.0 mg and 42.5 and 50.0 mg

for parasites with and without the 24 hour isolation interval, respectively.

This suggests that parasites presented with large hosts under these condi-

tions may be utilizing the maximum number of ovarian eggs suitable for

oviposition. Clutch size is somewhat smaller on NOW in the largest weight

classes for parasites which were not given 24 hour isolation intervals between

444 Contrib. Amer. Ent. Inst., vol. 20, 1983

hosts. This would be expected if egg utilization is at a maximum with large

hosts, as less time would be available for egg maturation between reproduc-

tive episodes.

6. Egg Production and Position: In addition to analyzing egg produc-

tion and host size, we examined the position of eggs with regard to segment

number and aspect (dorsal, lateral, ventral) of the host's body. Data pre-

sented in Table 4 were collected from the detailed tests on NOW (n = 26)

and PBW (n = 20) (both with 24 hour isolation interval) described above.

Data collected from NOW and PBW were maintained as distinct. From

Table 4 it is evident that there is a strong preference for the dorsal aspect

of the host's body, followed by a moderate preference for the lateral as-

pect of the host's body, while the ventral aspect of the host's body is

strongly rejected. The trend was consistent for PBW and NOW. Goniozus

legneri prefers to oviposit on segment 5-9 when attacking NOW and seg-

ments 4-9 when attacking PBW. Both trends were seen on the dorsal and

lateral aspects of the host's body for NOW and PBW. It was also noted for

the ventral surface of segments 5-9 on NOW and 5-8 on PBW.

This data was compared to similar data collected on two related

species, G. gordhi Evans (attacking Deoclona yuccasella Busck) (Gelichi-

idae) and G. aethops Evans (attacking PBW). Interestingly, G. gordhi

prefers to oviposit on the dorsal aspect of segments 6-9, has a moderate

preference to the lateral aspect of segments 6-9, and is reluctant to

Oviposit on the venter. This data is consistent with the findings for G.

legneri, but is in sharp contrast with the findings for G. aethops , which

demonstrates a strong preference for the lateral aspect of segments 5 -9,

but strongly rejects the dorsal and ventral aspects of the host's body.

7. Immature Mortality: Mortality of larvae and pupae were compared

for the 26 females attacking NOW and the 20 females attacking PBW

(Table 5). Mean egg production (at 25 + 2°C, 50% RH, LD 14:10) per fe-

male per lifetime was 288.00 + 33.91 on NOW, and 294.75 + 47.74 on PBW.

These results were not significantly different. These females produced

272.08 + 32.09 larvae on NOW and 284.50 + 46.24 larvae on PBW (not

significantly different). Thus egg-larval mortality was 15.54 + 6.21 individ-

uals on NOW, and 10.70 + 3.74 individuals on PBW. Pupal mortality was

6.20 +/ 3.80 individuals on NOW and 8.70 + 11.25 individuals on PBW.

8. Life Tables: Life tables were constructed for 25 female c.

legneri on NOW (one female was lost during the experiment) and the 21

female G. legneri on PBW. These parasites were provided with 24 hour

isolation intervals between ovipositions and hosts were chosen at random

from host cultures. Age (x) was counted in days; survivorship (1,) and age-

specific fecundity (mx) values were calculated by the methods described

above.

The life table incorporating age (x), survivorship (1,) and fecundity

(m,) values for G. legneri on NOW is presented as Table 6. Only the

values through the age of last female egg production (x = 105 days) are

shown. Table 7 is a life table for G. legneri on PBW and is presented in

a similar manner (age of last female egg production is x = 119 days).

Gordh, Woolley & Medved: Biology of Goniozus 445

From the life tables, demographic statistics were estimated using the

methods discussed in Birch (1948). Net reproductive rates (Rg) were cal-

culated by the formula:

Rt eo meh (1)

Generation times AT. 5) were calculated using the formula for cohort genera-

tion time:

be ie x1 m /Ro (2)

Two population growth rate parameters were calculated. Capacities for in-

crease (r) were derived using:

ee ee ee (3)

Intrinsic rates of increase a were estimated by iterative solutions of the

equation:

‘a a (4)

By this method, successive values of r_ are tried until the desired degiee of

accuracy is achieved. The capacity for increase (rc) is often used as an

approximation of the intrinsic rate of increase (r_), but is often discussed

by Laughlin (1965), May (1976), and others, both Statistics have relevance to

discussions of population growth phenomena. The capacity for increase (r_)

can be thought of as the growth rate of populations with non-overlapping

generations, or as the actual growth rate of a population founded by a single

cohort of females until a stable age distribution is obtained (i.e., for the

first several generations). Once a stable age distribution is obtained, a

population grows at the intrinsic rate of increase (r,,) in an unlimited en-

vironment. The capacity for increase (r_) is typically less than r,, and

Laughlin (1965) states that the differencé is usually small (5% or less)

while May (1976) provides a formula to estimate the relative error in esti-

mating the relative error in estimating fe, by t-

The demographic statistics we calculated for G. legneri are presented

in Table 8. Net reproductive rates (Rg) for this parasite on PBW and NOW

are 160.2 and 145.1 females per female per lifetime, respectively. Values for

generation time (T,) and capacity for increase (ro) were 51.5 days and

0.0985 females per female per day on PBW and 45.1 days and 0.110 females

per female per day on NOW. The intrinsic rates of increase (r,,) were

0.157 females per female per day on PBW and 0.162 females per female per

day on NOW. On both hosts, the differences between 1,, and ro were sub-

stantial: rp, exceeded ro by 37.3% on PBW and by 32.1% on NOW. The

differences are due to the large Ro values and, possibly, to large variances

in the 1m, distributions (see May [1976] for a discussion of this problem).

446 Contrib. Amer. Ent. Inst., vol. 20, 1983

We wished to compare the demographic statistics for G. legneri with

those produced for G. emigratus on NOW by Gordh and Hawkins (1981).

However, the value given for r,, in that paper was calculated using formula

(3) and is therefore better referred to as r~. We have included this sta-

tistic (as ro and those for net reproductive rate (Ro) and cohort

generation time (T_) in Table 8. In order to compare intrinsic rates of in-

crease between the 2 species of Goniozus, we have used the life table data

presented in Gordh and Hawkins (1981) to calculate 1, for G. legneri using

equation (4). This is also included in Table 8. A similar trend for the

"approximation" statistic (r_) versus the "accurate" statistic (rm) was noted

for G. emigratus on NOW: “r_ was 0.178 and exceeded r_ (0.131) by 26.4%.

These empirical results indicate that the choice of one méthod or another to

estimate population growth rates is not necessarily trivial as substantially

different results may be obtained. The differences between rm and f¢

values for these Goniozus spp. are only slightly greater than the differences

predicted by applying May's (1976) formula to our life table data.

The life table for G. emigratus on NOW (Gordh and Hawkins 1981)

was obtained using an experimental protocol virtually identical to that used

for G. legneri as described above. We note one difference: G. emigratus

females were maintained at 26.7 + 1°C and G. legneri was maintained at

25. + 2°C on both hosts in our experiments. Nevertheless, values for demo-

graphic statistics computed by equivalent methods should be directly

comparable between the two species reared on NOW, and between hosts for

G. legneri. The value for r_ for G. emigratus on NOW is substantially

greater than that obtained for G. legneri on the same host, 0.178 versus

0.162 females per female per day, respectively, even though the net repro-

ductive rate for G. emigratus on NOW (R, = 128.0 females per female per

lifetime) is substantially less than that for G. legneri on NOW (Ro =

145.1 females per female per lifetime). This effect is due to the shorter

generation time for G. emigratus on NOW (T = 27.3 days) versus

legneri on NOW (T = 30.7 days), and may be due, in part, to the temper-

ature difference noted above. Similarly, G. legneri has a slightly higher

ron NOW than on PBW (0.162 and 0.157 females per female per day,

respectively) despite the lower R, obtained for this parasite on NOW

(145.1 females per female per lifetime versus R, = 160.2 females per female

per lifetime on PBW). Again, the slightly higher r,. on NOW is due to the

shorter generation time for G. legneri on this host: 30.7 days versus 32.4

days on PBW.

The net reproductive rates (R,) for G. legneri on PBW and NOW ob-

tained by analyses of life tables were very close to the mean number of

female progeny on PBW and NOW (169.55 + 28.80 and 149.21 + 17.62 females

per female per lifetime, respectively) obtained by direct averaging of the

data. One possible reason for the greater total production of female prog-

eny by G. legneri on PBW is discussed above under 2. Parasite Host-

Attack. That is, the greater number of males in the original broods of

G. legneri on PBW resulted in more sperm per female per insemination.

Gordh, Woolley & Medved: Biology of Goniozus 447

The difference cannot be due to host size (larger hosts supporting larger

broods with more females) as PBW hosts tended to be smaller than NOW

hosts in these experiments (see Table 3). The shorter generation time

(T) for G. legneri on NOW is difficult to account for, but it may be re-

lated to host size or host quality.

DISCUSSION

In recent years there has been considerable interest in the study of

bethylid biology and the potential of bethylids in biological control of vari-

ous agricultural and urban pests. Much of this work has been reviewed else-

where and need not be repeated here (see Gordh 1976, Gordh and Evans

1976, Gordh and Hawkins 1981, and papers cited therein).

Studies have shown that several species are of value to biological con-

trol, and G. legneri seems to be another useful species in attacking

Microlepidoptera. The fossorial habits of bethylids, their longevity, and high

reproductive potential would be desirable attributes for biological control

agents attacking Lepidoptera and Coleoptera larvae, especially in concealed

situations. Although not host specific, they do demonstrate strong prefer-

ence at the family or superfamily level.

Although considered among the most primitive of aculeate Hymenop-

tera, bethylids display a highly complex biology. Within the genus

Goniozus, several species are known to regulate the number of eggs

deposited based on an estimate of host size. The modalities responsible for

this phenomenon are not known, but they are currently under investigation.

Regulation of sex ratio based on egg clutch size is also apparently wide-

spread in Goniozus, and will perhaps be found in other bethylids as well.

Further research on other bethylids will prove exceptionally rewarding in

understanding the evolution of the aculeate Hymenoptera.

The morphology of the immature stages of G. legneri do not differ

from other species of Goniozus (cf. G. columbianus, G. japonicus, G.

emigratus, G. gordhi, G. aethops, G. claripennis Foerster, G. nigrifemur

[Ashmead]). There are some indications that the manner in which pupal

pigmentation is formed differs among species, and may be of some tax-

onomic value (Gordh 1976, Gordh and Evans 1976). Unfortunately this

aspect of immature development has not been critically examined for most

species.

The longevity of G. legneri is substantially longer than many other

species of Goniozus which have been studied. For instance, Gordh and

Evans (1976) found that G. aethops lived 13.35 + 8.48 days, Gordh and

Hawkins (1981) found that G. emigratus lived 52.60 + 7.30 days when pro-

vided hosts and 37.10 + 12.40 days when deprived of hosts, and Gordh

(1976) found that G. gordhi females lived 62.43 + 8.48 days when provided

hosts. Thus it appears that there is considerable variability in the longe-

vity of Goniozus species.

There is similarly some variability in the number, size and position of

eggs of Goniozus. Gordh (1976) reported that G. gordhi deposited 153.78 +

48.59 eggs during the course of a female's lifetime, Gordh and Evans (1976)

448 Contrib. Amer. Ent. Inst., vol. 20, 1983

reported that G. gordhi deposited 153.78 + 48.59 eggs during the course of

a female's lifetime, Gordh and Evans (1976) reported that G. aethops de-

posited 16.23 + 8.18 eggs during the course of its lifetime, and Gordh and

Hawkins (1981) found that G. emigratus deposited 232.9 + 91.5 eggs per life-

time under conditions similar to those provided G. legneri.

Goniozus legneri has a strong ovipositional preference for the dorsal

aspect of the host's body. This site preference is also seen in G. gordhi

(Gordh 1976), G. cellularis (Say) (Gordh and Evans 1976), G. nigrifemur

(Gordh, unpubl.), and G. emigratus (Gordh and Hawkins 1981). Other

species (G. aethops, Gordh and Evans 1976, G. nephantidis, Antony and

Kurian 1960) prefer the lateral aspect of the host's body while Cherian and

Israel (1942) indicated that G. indicus prefers the ventral aspect of the

host's body as an ovipositional site.

The size and shape of Goniozus eggs can vary also. Most species have

the typical sausage-shaped (hymenopteriform) eggs which are deposited

longitudinally along the host's body, but some species have "football" shaped

eggs which are deposited intersegmentally within the integumental folds

(G. aethops, Gordh and Evans, 1976). The thickness of the chorion also ap-

pears to be related to the position of the egg and the extent of paralysis.

Permanently paralyzed hosts with eggs deposited on segments have relatively

thin chorions; eggs which are deposited on hosts which recover from paraly-

sis and resume activity tend to have football shaped eggs which have thick

chorions and are deposited intersegmentally.

In all species of Goniozus studied, there appears to be a normal dis-

tribution of eggs over the integument of the host. That is, all species pre-

fer to oviposit on the middle segments of the host's body, irrespective of

the aspects which it prefers (i.e., dorsal, lateral, ventral) and progressively

fewer eggs are deposited on segments toward the head or anus. The purpose

for this behavior is not clear. Hosts paralyzed permanently can exhibit some

lateral movement. Considering the thin chorion of most eggs, and the pos-

sibility of dislodgement, and subsequent mortality, then it is unlikely that

the middle segments of the body would be a suitable ovipositional site be-

cause in lateral flexure, these segments are exposed to the most deforma-

tion. Alternatively, it appears that maximum haemolymph flow is the most

important consideration in survival of the parasite. At least, hosts which

have been killed by the female parasite and oviposited upon yield no para-

site progeny. Given the shape of the host larva, it seems probable that

maximum haemolymph flow (volumetrically) is located along the middle

segments of the host's body. Thus we infer that the reason the middle

segments are preferred is due to haemolymph flow.

Courtship in G. legneri is not different from that noted for c.

gordhi, Gy. aethopes, Ga.emigratus,.G.nephantidis.).C; montanus., OF. GC.

claripennis. Indeed, in all species of Goniozus and related genera court-

ship is overtly simple. In all species of Goniozus it appears that the male

chews into the female's cocoon. Thus sib mating in broods producing males

and females appears to be the rule.

The host attack by G. legneri does not differ from other species of

the genus. Invariably the female stings her host in the ventral nerve cord

near the head. The only noted exception to this site specificity is a report

Gordh, Woolley & Medved: Biology of Goniozus 449

by Bridwell (1919) in which he remarked that in G. emigratus the parasite

stung its host in three places ~ the head region, the middle of the body,

and near the anus. In a subsequent study (Gordh and Hawkins 1981) this

behavior was not observed in G. emigratus. We suspect that Bridwell's

comments were based on a very limited number of observations, and that in

fact what he noted was a female parasite initially mounting the host at the

posterior end of its body and that in the ensuing struggle, the parasite

shifted to the head. This sort of behavior has been observed in G. emi-

gratus, G. aethops, G. legneri, and G. gordhi.

There does appear to be some variability in the effect of venom in

Goniozus. In most species the paralysis is permanent, but in G. japonicus

(Iwata 1961, Kishitani 1961), Gc. rugosus (Samad 1973), and G. claripennis

(Voukassovitch 1924), the host regains the ability to walk after the parasite

has oviposited. Although the reports on regaining locomotion are not clear

on this point, it is suspected that any species of Goniozus which temporar-

ily paralyzes its host probably has small eggs with thick chorions, and de-

posits its eggs in the intersegmental folds of the host's integument. Other-

wise, high egg mortality could be suspected.

A post-adult emergence period during which the parasite is incapable

of oviposition seems common in Goniozus, but its duration is highly variable.

Gerling (1979) found that Goniozus sp. had a preovipositional period of up to

nine days, and Iwata (1961) found that G. japonicus has a preovipositional

period of about seven days. Most other species of Goniozus which have

been studied have a preovipositional period of 3-5 days.

Demographic statistics such as we have calculated for G. legneri may

eventually prove to be useful to biological control workers in evaluating the

potential of natural enemies or in analyzing performance in particular habi -

tats. Certainly, many other attributes of natural enemies are responsible for

success or failure in a given program. These statistics do provide a good

measure of reproductive potential in the laboratory. The choice of which

population growth statistic to use is best determined by the context of par-

ticular comparisons. For example, r may be close to the actual growth

rate of a population of natural enemies soon after a release has been made.

If hosts are abundant and competition effects negligible, the growth rate

may increase for several generations and approach rp, if a stable age dis-

tribution of parasites is obtained. The me values which we have calculated

for G. legneri on PBW and NOW (0.157 and 0. 162) and for G. emigratus

on NOW (0.178) are rather high, even for insects (see Connell and

Scheiring (1982) for a recent tabulation of r_ values for various insects).

This indicates that these species of Goniozus are capable of rapid increases

in population size under favorable conditions.

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Szeged 16: 163-165.

Moczar, L. 1971. Pycnomesitius New Genus from Bethylidae (Hymenop-

tera). Acta Biol. Szeged 17: 167-169.

Moczar, L. 1974. On another new species of the genus Metrionotus

Moczar. Acta Biol. Szeged 20: 173-177.

Méczar, L. 1975. Two new species and a key for the males of the genus

Heterocoelia Dahlbom (Hymenoptera: Bethylidae, Mesitinae). Acta

Zool. Acad. Sci. Hung. 21: 109-114.

Moczar, L. 1976a. New Sulcomesitius species from Borneo. Acta Biol.

Szeged 22: 97-105.

Moéczar, L. 1976b. New species of Sulcomesitius Méczdr from Laos

(Hymenoptera, Bethylidae). Ann. Hist. Nat. Mus. Nat. Hung. 68: 279-

286.

Moczar, L. 1978. Two new species of Mesitinae from Egypt (Hymenop-

tera: Bethylidae). Acta Biol. Szeged 24: 139-142.

Moczar, L. 1981. New Sulcomesitius and Heterocoelia species from SE

Asia (Hymenoptera: Bethylidae). Acta Zool. Acad. Sci. Hung. 27: 355-

a073

Nagy, C. 1966. Neue Bethyliden-arten fur die fauna Rumaniens (Hymenop-

tera). Trav. Mus. Hist. Nat..6: 165-167.

Nagy, C. 1967. Contributions a la connaissance des hyménopteres Bethyli-

dae de la faune roumaine. Folia. Entomol. Hung. N.S. 20: 87-94.

Nagy, C. 1968a. Les femelles de Laelius anthrenivorus Trani (Hym.:

Bethylidae) attaquent I'homme. Riv. Parasit. 29: 71-74.

Nagy, C. 1968b. La presence du genre Holepyris Kieff. (Hym.: Bethyli-

dae) dans la faune de la Roumanie. Jassy. Univ. Anal. Stiint. Sect.

2A, 14: 409-410.

Nagy, C. 1968c. A new record of Mesitinae (Hymenoptera, Bethylidae).

Mem. Ital. Ent. Soc. 47: 168-176.

Nagy, C. 1969. Further records on Mesitinae (Hymenoptera: Bethylidae).

Acta Zool. Acad. Sci. Hung. 16: 205-208.

Nagy, C. 1970a. Uber die in Europa vorkommenden mikropterischen

Bethyliden ~ Arten. Nachr. Bayer. Entomol. 18: 61-64.

Gordh, Woolley & Medved: Biology of Goniozus 453

Nagy, C. 1970b. Contribution toward a revision of the European species of

the genus Epyrus Westw. (Hym., Bethylidae). Mitt. Zool. Mus. Inst.

Univ. Berlin 46: 265-272.

Nagy, C. 1972. Taxonomic remarks on Mesitinae (Hymenoptera: Bethylidae).

Mem. Ital. Ent. Soc. 51: 5-18.

Nagy, C. 1974. A new bethylid subfamily allied to Protopristocerinae

(Hymenoptera, Bethylidae). Bol. Ital. Ent. Soc. 106: 126-130.

Nagy, C. 1976. Bethylidae (Hymenoptera) parasitizing orchard caterpillars.

Rev. Roum. Biol. Ser. Biol. Anim. 21: 103-108.

Remadevi, O. K. etal. 1978. Oviposition behavior of Perisierola

nephantidis Muesebeck (Bethylidae: Hymenoptera), a larval parasite of

Nephantidis serinopa Meyrick (Xylorictidae: Lepidoptera). Entomology

3: 303-305.

Samad, K. 1973. Systematics and biology of Goniozus rugosus ~ new

species (Bethylidae: Hymenoptera) a larval parasite of microlepidoptera

in Pakistan. Agric. Pakistan 24: 195-200.

Venkatraman, T. V. and Chacko, M. J. 1961. Some factors influencing the

efficiency of Goniozus marasmi Kurian, a parasite of the maize and

jowar leaf roller. Ind. Acad. Sci. Proc. 53: 275-283.

Voukassovitch, M. 1924. Sur la biologie de Goniozus claripennis FoOrst.

parasite d'Oenophthira pilleriana Schiff. Bul. Soc. d'Hist. Natur.

Toulouse 52: 225-246.

Yamada, Y. 1955. Studies on the natural enemy of the woolen pest,

Anthrenus verbaci Linne (Allepyris microneurus Kieffer) (Hymenop-

tera, Bethylidae). Mushi 28: 13-29.

454 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table 1. Mean and standard deviation for sting to oviposition, egg deposition

to eclosion, larval stage duration, pupal stage duration, and egg to

adult duration for broods producing only males and broods producing

males and females on NOW and PBW.

Host

Interval PBW NOW

Sting - Oviposition

he 1.40 + 0.59 (n = 429) 1.19 + 0.47 (nm = 437)

oo 1.96 2 1.07; 40. = 71) 1.820% 1216: (n-= .TS9)

Oviposition - Eclosion

JF y 1.44 + 0.18 (n = 428) 1.25 + 0.27 (mn = 436)

go 1.46 + 0.13 (n = 71) 1.23 + 0.27 (mn = 155)

Larva - Cocoon

J? 3.61 + 0.26 (n = 428) 3075 2°0235. Cae e430)

Jo 3.56 & 0223 (a= /71) oi 60°8 0321 {nH 2155)

Cocoon —- Adult

J? 8.93 £0.57 (a: = 428) 9.26 + 0.53 (n= 430)

Jo 8.96 £ 0,38 @ = 71) B6a 3° O000. ae 155)

Egg -— Adult

ed 13-98. 2 °O553) (i427) 14.26 + 0.97 (n = 430)

oo 14.00 20.47 ‘(a =. 72) £3.43 20.46 (na = 155)

Gordh, Woolley & Medved: Biology of Goniozus 455

Table 2. Summary (joint frequency distribution) of data for complete broods

for first 12 broods for each female parasite. (Female parasites

began exhausting sperm supply on 13th brood.)

NUMBER OF MALES

Brood size 0 1 Z 3 4 Total

5 3 3

6 4, 4

i 8 8

8 10 2 ta

9 5 8 13

10 ai 1 22

11 29 3 ae

ee 33 8 41

13 LS 14 a9

14 6 9 15

LS 9 9

16 2 2

17 0

18 0

19 . 0

20 1 ;

Total 0 30 114 46 1 191

1983

Contrib. Amer. Ent. Inst., vol. 20,

456

se sire 0 3 6°91 o8 sure ee 0 OG 4940

€ Ove 9¢ sa 0 ish a 0) 0°0S—-S° LY

c Bet Se él 8° ST Goce = = 0 S L790 S77

9 Gel SL 02 ae LLe € ot 87 O°S7-S ZY

OT oe! ITT 8c OSs 617 L 1 Ss 90T 29-0 OF

6T Cot TSZ 09 oes cE8 TT V1 £ST O Ors 2

02 7 C1 872 T6 £° at 9STI L de co | 68 S 14-0 SE

02 eee | 677 80T Ae is 6TET 9S O° Et EoL Go Se-S et

es O21 L6E 60T 61 7671 cEl s'il 9ST £*ct-0 OE

eT 16 60T GV Pee: 667 98 ae a 6201 “£270 Sd

C O°OT Od 9T 7°6 OST BE £°Ot SOY OSes £2

ee — 0 8 7°83 Se a6 eh 707 G°ée-0°02

19> aig 0 ag 0 9 G°L GY O¢ tepun

u qsoy aod S830 t- 3soy zod s38o u 3soy aod s330 (3u) 343ToOM

s33q TeI207, s33q TeqoO] s33q Tero] 3sS0H

(TBATeAUE ou) (TBALSeIUT anoy 47) (TB@ATeAWUTE ANoYyY 47)

9 = U) MON (892 = 1) MON (s0Z = &) Maa

| 1S0H

*peaAresqo sjsoy Jo Jequnu = U ‘UPA YORe AO *SiSsOoY UseM}IEeq TeATEUT ANOY 47 ANOYITM pue UATM

‘,y3TemM Jsoy FO UoTIOUNZ e se Mgq pue MON UO Jsoy zed s388e Jo rzequnu ueoew pue uoTIONpoad 33a TeIoT ‘*¢€ eTqe]

Gordh, Woolley & Medved: Biology of Goniozus

Table 4. Cumulative number of eggs produced per segment and egg position by G. legneri

attacking NOW (n = 26 female parasites) and PBW (n = 20 female parasites).

Data compared with similar data collected for G. gordhi Evans, G. aethops

Segment of host's body

Aspect Z, 3 4 5 6 7 8 9 10 11

G. LEGNERI (NOW)

Dorsal 5 8 58 258 9621; 1454 1456 699 118 0

Lateral 0 3 10 85 431 Hie 649 292 40 )

Ventral 0 1 0 ie 38 72 54 15 0 0

G. LEGNERI (PBW)

Dorsal 14 67 32 678 1021 1067 924 426 76 2

Lateral 1 11 57 144 278 312 245 117 25 2

Ventral e 8 8 va KY 25 a7 5 v4 1

G. GORDHI (Deoclona yuccasella Busck)

Dorsal 0 5 iz 56 120 193 202 153 3 0

Lateral 0 a 9 21 39 67 42 92 14 5

Ventral 0 i 2 5 4 2 6 3 2 0

G. AETHOPS (PBW)

Dorsal 0 0 2 2 3 6 8 4 0 0

Lateral 4, 6 26 fg 72 iG 118 81 22 1

Ventral 0 0 2 1 0 5 0 0 0 0

458

Contrib. Amer. Ent. Inst., vol. 20,

1983

Table 5. Immature mortality of G. legneri on PBW and NOW. Each figure

Eggs

329

207

358

326

317

338

295

287

274

Sh7

318

253

299

263

278

288

PW 5

263

eae:

267

269

209

226

280

288

288.00

ei Be ee

represents the total lifetime production of one female parasite.

NOW

Larvae

301

197

337

308

285

331

284

272

255.

307

302

234

271

251

271

270

265

253

301

257

263

250

281

205

272

261

212-

(28)

(10)

(21)

(18)

(22)

(17)

(11)

(id)

(19)

(10)

(16)

(19)

(22)

(12)

(7)

(18)

(8)

(10)

(14)

(10)

(4)

(19)

(18)

(21)

(8)

eee)

Adults

293

L941

330

298

283

31it

282

272

246

304

298

230

260

243

263

264

256

254

297

248

260

244

280

LoD

256

200

266.

£32 e

(8)

(6)

(2)

(10)

(2)

(10)

(2)

(0)

(9)

(3)

(4)

(11)

(8)

(6)

(9)

(2)

(4)

(9)

(3)

(6)

(1)

(10)

(16)

(2)

PBW

Larvae

222

308

323

319

219

178

o2i

261

328

332

27%

250

322

359

286

245

286

322

265

261

284;

+46.

(5)

(11)

(7)

(6)

(11)

(9)

(13)

(12)

(13)

(17)

(9)

(18)

(9)

(14)

(7)

(9)

(7)

Figures.in parenthesis refer to mortality in that stage.

Adults

281

296

318

272

210

Wis

aug

254

324

268

249

317

356

282

Z19

276

319

263

258

yay go

+46.

(4)

(18)

(5)

(47)

(9)

(2)

(7)

(4)

(8)

(3)

(i)

(8)

(3)

(4)

(30)

(10)

(3)

(2)

(3)

Gordh, Woolley & Medved: Biology of Goniozus 459

Table 6. Life table for G. legneri on NOW. Developmental stage, age (x)

in days, survivorship (1,)> and age-specific fecundity (m,)>

n = 25 #

Developmental Age Survivorship Fecundity

Stage x 1. m

eggs 0 1.000 0

lst instar larvae 1 945 0

larvae to adult emergence Zo NS 924 0

mating and isolation period 14 - 15 ~924 0

hosts available 16 924 0

17 924 0

18 924 i 200

19 924 6.620

20 924 480

21 924 0

22 924 9.480

23 ~924 0

24 924 4.760

Zo 924 3.600

26 924 760

27 924 4.400

28 924 3.560

29 924 2.840

30 924 0

31 924 5.600

32 ~924 2.640

33 ~924 . 600

460 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table 6 (continued).

Age Survivorship Fecundity Age Survivorship Fecundity

x 1. ma x 1. Mm

34 924 7e720 58 924 960

35 924 440 29 924 4.080

36 924 640 60 924 2.880

37 924 2.800 61 924 LV 360

38 924 4.760 62 924 2.760

a8 924 1.000 63 924 2.840

40 924 2.680 64 924 1.680

41 924 4.760 65 924 23900

42 924 2.080 66 924 1.640

43 924 2.000 67 924 3.000

44 924 4.160 68 924 1.320

45 924 evo20 69 924 Le720

46 924 3.400 70 924 1.880

47 924 2:20 sgt 924 1.400

48 924 a7 20 as 924 1.640

49 924 26760 Ls 924 eae

50 924 1.160 74 924 840

ep 924 3.240 ee, 924 840

oe 924 hae 4 76 924 960

53 924 3.640 77 924 1.160

54 924 1.280 78 924 080

55 924 2.080 #9 924 480

56 924 5.960 80 924 .880

57 924 1.800 81 924 «POO

Gordh, Woolley & Medved: Biology of Goniozus 461

Table 6 (continued).

Age Survivorship Fecundity Age Survivorship Fecundity

x 1. m x 1. i ma

82 924 - 240 104 “O40 0

83 924 040 105 «ek3 -046

84 924 - 240 106 813 0

85 924 120

86 924 - 160

87 924 040

88 924 0

89 924 - 080

90 924 . 160

91 924 0

92 ~924 - 040

93 ~ 924 0

94 ~924 -040

95 Ae . 080

96 924 0

97 887 042

98 887 0

99 ~887 0

100 887 0

101 887 0

102 887 0

103 - 887 0

462 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table 7. Life table for G. legneri on NOW. Developmental stage, age (x)

in days, survivorship (1s and age-specific fecundity (m) >

n= 21 ¢

Developmental Age Survivorship Fecundity

Stage x 1. ma

eggs 0 1.0000 0

Ist instar larvae a +9637 0

larvae to adult emergence Z\= 13 9345 0

mating and isolation period 14:- 15 ~ 9345 0

hosts available 16 ~9345 0

17 -9345 0

18 ~ 9345 2.143

19 - 9345 b+239

20 -9345 333

at 9345 25095

22 ~9345 3.560

23 ~ 9345 2.047

24 ~ 9345 1.905

25 ~ 9345 3.471

26 ~ 9345 1.286

27 ~9345 15952

28 ~9345 3.505

29 9345 2.762

30 ~ 9345 2.286

31 9345 2.762

32 -9345 2.143

30 ~9345 2.492

Gordh, Woolley & Medved: Biology of Goniozus 463

Table 7 (continued).

Age Survivorship Fecundity Age Survivorship Fecundity

x 1. m x 1. ma

34 9345 et be 58 9345 2.143

Ee) 9345 2.714 59 9345 1.286

36 9345 eases 60 9345 5.238

37 9345 Ba dOe 61 9345 Ze 359

38 9345 L.190 62 9345 1.810

39 9345 3.524 63 9345 1.381

40 9345 2.762 64 9345 4.0695

41 9345 714 65 9345 2.905

42 9345 3.619 66 9345 1.429

43 9345 3.476 67 9345 1.476

44 9345 476 68 9345 2.857

45 9345 4.048 69 9345 2.905

46 9345 4.095 70 9345 810

47 9345 1.429 71 9345 1.524

48 9345 2+399 re 9345 2.414

49 9345 4.571 13 9345 ya ee |

50 9345 1.714 74 8900 700

5} 9345 2.143 ih) 8900 1.450

52 9345 3.286 76 8900 2.200

ao 9345 2.714 a 8900 1.400

54 9345 952 78 8900 850

55 9345 3.667 19 8900 3.100

56 - 9345 3.524 80 - 8900 1330

on 9345 oraeL 81 . 8900 1.500

464 Contrib. Amer. Ent. Inst., vol. 20, 1983

Table 7 (continued).

Age Survivorship Fecundity Age Survivorship Fecundity

x 1. m x 1. m

82 -8900 - 700 | 105 - 6230 0

83 -8900 i320 106 - 5340 .083

84 -8455 1.368 107 - 5340 0

85 ~8455 947 108 - 5340 +333

86 8455 Lezes 109 - 5340 2290

87 ~8455 <ar9 110 -5340 0

88 8455 684 a 4450 . 100

89 -8455 1.632 112 4450 0

90 8455 - 368 113 -4450 .700

OL 8455 «158 114 ~4450 0

92 -8455 947 115 -4450 0

93 -8010 444 116 -4450 0

94 1965 e765 hig - 4005 eae

95 7565 . 706 118 Ms 1B a 0

96 ay Be obs oou ce, Pec . 286

ay Me pel ok 059 120 - 2670 0

98 »7565 824

99 -7120 1.000

100 6675 woo

101 -6675 0

102 -6675 467

103 -6230 1,500

104 - 6230 214

Gordh, Woolley & Medved: Biology of Goniozus 465

Table 8. Demographic statistics for G. legneri on PBW (n = 21), G. legneri

on NOW (n = 25), and G. emigratus on NOW. Statistics for G.

emigratus taken from (*) or calculated from (**) Gordh and

Hawkins (1981).

G. legneri G.. legneri G. emigratus

Statistic PBW NOW NOW

Ls Ro 160.2 | 145.1 128.6

2 qT. 2133 45.1 Be uth

3. ry .0985 -110 .131

4. th ~157 - 162 -178

1. Net Reproductive Rate, females per females per lifetime, Roy = tL om

2. Cohort Generation Time, in days, T = 2 xl m/R

c x sO

3. Capacity for Increase, females per female per day, to°% In Ro/T.

oo ee)

4. Intrinsic Rate of Increase, females per female per day, as x21 Ime

466 Contrib. Amer. Ent. Inst., vol. 20, 1983

ane)

100

-. oa

3 Be

<1 . 0

NOW MATED UNMATED

PBW - UNMATED J

MATED Y W/HOSTS NO HOSTS

Fig. 1. Mean and variance of longivity of G. legneri comparing mated

parasites provided hosts (NOW, PBW) (n = 50 each), and mated and unmated

females deprived of hosts (n = 50each), and unmated males (n = 30).

Gordh, Woolley & Medved: Biology of Goniozus 467

o PARALYZED AND PARASITIZED

e@ PARALYZED

can POs

3 sol

ot eect Hat |

ee +}

60 ? HT | i F

ent

D 30 ;

Fig. 2. Mean and variance of pulse rate of NOW which were paralyzed

(dots) and which were paralyzed and parasitized (circles) by G. legneri during

their lifetime (n = 10 for each sample).

468 Contrib. Amer. Ent. Inst., vol. 20, 1983

DORSAL VENTRAL DORSAL VENTRAL

eo: HRS 48 HRS

DORSAL VENTRAL DORSAL VENTRAL

TS et ve HRS

Fig. 3. Dorsal and ventral aspects of 24, 48, 72, and 96 hour old

pupae of G. legneri, illustrating the sequence of pupal pigmentation.

NEW TAXA DESCRIBED IN THE VOLUME

Mattias Idar

Hadrodactylus townesi, n. sp. (Ichneumonidae)

W. R. M. Mason

Khoikhoiinae, n. subfamily (Braconidae)

Sania, n. genus mR

Sania henryi, n. sp.

S. marjoriae, n. sp.

S. capensis, n. sp.

Khoikhoia, n. genus ms

K. townesi, n. sp.

K. solata, n. sp.

K. semiadusta, n. sp.

Lars Huggert and Lubomir Masner

Labidopria noctivaga, n. sp. (Diapriidae)

Townesella, n. genus

T. marjoriae, n. sp.

Plagiopria, n. genus

P. passerai, n. sp.

P. besucheti, n. sp.

P. huberi, n. sp.

Trichopria myrmicae, n. sp.

C. M. Yoshimoto

Henryana, n. genus (Eulophidae)

H. magnifica, n. sp.

M. J. Sharkey

Marjoriella, n. genus (Braconidae: Agathidinae)

M. ancha, n. sp.

_M. beni, n. sp.

M. henryi, n. sp.

M. townesorum, Nn. sp.

D. R. Kasparyan

Lygurus, n. genus (Ichneumonidae: Phrudinae)

L. townesi, n. sp.

P. M. Sanborne

Sesioplex brunipalpus, n. sp. (lchneumonidae)

S. cribus, n. sp.

469

Page

116

ive

172

470 Contrib. Amer. Ent. Inst., vol. 20, 1983

Virendra Gupta

Dyspetes townesi, n. sp. (Ichneumonidae: Tryphonini) 183

D. flavus, n. sp. 183

D. indicus, n. sp. 186

Testsusaburo Tachikawa

Astichus townesi, n. sp. (Eulophidae) 189

|. D.Gauld and G. A. Holloway

Tryonocryptus, n. genus (Ichneumonidae: Endaseini) 192

T. nigridorsalis, n. sp. 193

T. gigas, n. sp. 194

‘T. amicus, n. sp. 195

L. Moczar

Sulcomesitius townesianus, n. sp. (Bethylidae) 198

Reijo Jussila

Atractodes townesi, n. name (ichneumonidae) 202

A. fittoni, n. name 202

Charles C. Porter

Trachysphyrus townesi, n. sp. (Ichneumonidae) 207

Karl-Johan Hedaqvist

Karpinskiella townesi, n. sp. (Pteromalidae) tog I

Maximilian Fischer

Opius (Grimnirus) eutownesi, n. sp. (Braconidae: Opiinae) 223

O. (G.) ileifensis, n. sp. 225

O. (G.) nigeriensis, n. sp. 221

O. (G.) obuduensis, n. sp. 229

O. (G.) toromojaensis, n. sp. 231

O. (Frekius) nairobicus, n. sp. 235

J. Papp

Biosteres (Biosteres) townesi, n. sp. (Braconidae: Opiinae) 237

Opius (Apodesmia) marjorieae, n. sp. 239

Opius (Utetes) townesianus, n. sp. 241

Santosh Gupta

Prosthoporus townesi, n. sp. (Ichneumonidae: Gabuniini) 247

P. striatifrons, n. sp. 247

P. panamensis, n. sp. 248

P. marjoriae, n. sp. 249

P. nigrifemur, n. sp. 250

New Taxa

Michael G. Fitton

Bioblapsis henryi, n. sp. (Ichneumonidae: Diplazontinae)

Alexander P. Rasnitsyn

Praeichneumonidae, new family (Fossil Ichneumonoidea)

Praeichneumon, n. genus

P. townesi, n. sp.

Eobracon , n. genus (Braconidae)

E. inopinatus, n. sp.

Robert A. Wharton

Senwot, n. genus (Braconidae: Alysiini)

S. africanus, n. sp.

C. van Achterberg

Brulleiini, n. tribe (Braconidae)

Parabrulleia, n. genus

Brulleia nipponensis, n. sp.

B. townesi, n. sp.

B. brunnea, n. sp.

B. nigra, n. sp.

B. tricolor, n. sp.

Paul M. Marsh

Megaloproctus platyantennus, n. sp. (Braconidae: Doryctinae)

M. schunckei, n. sp.

M. striatus, n. sp.

M. townesi, n. sp.

M. xanthus, n. sp.

Erich Diller

Herpestomus henrytownesi, n. sp. (|chneumonidae: Phaeogenini)

E. Haeselbarth and Conrad Loan

Townesilitus, n. genus (Braconidae: Euphorinae)

Vinalto Graf

Philodrymus townesi, n. sp. (Ichneumonidae: Anomaloninae)

Luis De Santis

Tetramesa adrianae, n. sp. (Eurytomidae)

Eurytoma cardoi, n. sp.

Mercetina entreriiana, n. sp.

Mimencyrtus, n. genus (Pteromalidae)

M. stipitatus, n. sp.

Horismenus seminiger, n. sp. (Eulophidae)

471

254

259

263

264

278

285

287

289

290

291

292

373

374

376

381

384

398

404

408

410

412

413

apie ‘

a vel i

seat

aD Pats is

5 yep A

if aiat

; Ki

n c he a }

Leal

yet

a |

= Se tt he

gg otter

Le,

‘ky

TIP er shay

rate the ipne fa

Mie = erm 3 Tune Lone

ttt

MP4

Ta Ea 8 heer 5

= FS ee te Goowe a Berg Seed mf 4 sen Sly Foe ae Sa

LS 2 - Nace Sas + Ser a Sip he ge er ae " cone as 4 Se -

Sof ee a 5: a el, POSE iS hi re’, yr ~ + — : - - , lew

fh Bop tie . 3 5 ber es ore Si - aah : s- ow ne x te ure Sy % < L te,

. eae ; = “~ 4 ar of eit = ie ee oe 1 pm a Hehe te wwe Bene ge

ce - ‘ 3 3 Rs Ser oe tea eens Horo Sy -: i a Sipe >Re Ep = Bm Ge

PPP vo ’ pe hme Gor frm ner SP : fm trees

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